Established in 1960 the Intergovernmental Oceanographic Commission (IOC) of UNESCO promotes
international cooperation and coordinates programmes in research, services and capacity building, in
order to learn more about the nature and resources of the ocean and coastal areas and to apply that
knowledge for the improvement of management, sustainable development, the protection of the
marine environment, and the decisionmaking processes of its 135 Member States.
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IOC TECHNICAL SERIES NO. 69, VOL.2
1
Biodiversity and distribution of the megafauna
Vol.2
ANNOTATED PHOTOGRAPHIC ATLAS OF THE ECHINODERMS
OF THE CLARION-CLIPPERTON FRACTURE ZONE
Dr Virginie Tilot
Membre correspondant de l’Académie des Sciences d’Outre-Mer
Attaché honoraire du Muséum national d’Histoire naturelle
Département des Milieux et peuplements aquatiques
55 rue Buffon, F-75005 Paris, France
With the support of the Government of Flanders (Belgium)
UNESCO
ENGLISH ATLAS PHOTO pages.indd 1
2006
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The designations employed and the presentation of the material in this publication do not imply the expression of any opinion whatsoever
on the part of the Secretariats of UNESCO and IOC concerning the legal status of any country or territory, or its authorities, or concerning
the delimitation of the frontiers of any country or territory.
For bibliographic purposes, this document should be cited as:
Tilot, Virginie. Biodiversity and distribution of the megafauna. Vol. 1: The polymetallic nodule ecosystem of the Eastern Equitorial Pacific
Ocean; Vol. 2: Annotated photographic atlas of the echinoderms of the Clarion-Clipperton fracture zone. Paris, UNESCO/IOC, 2006 (IOC
Technical Series, 69).
This publication has received financial support from the Government of Flanders (Belgium).
Published in 2006 by the United Nations Educational, Scientific and Cultural Organisation
7, Place de Fontenoy, 75352 Paris 07 SP
Page layout: Eric Loddé
Printed in the workshops of UNESCO
(c) UNESCO 2006
Printed in France
(SC-2006/WS/11vol.2)
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Table of Contents
Echinoderms of the Clarion-Clipperton fracture zone ......................2
Crinoids .......................................................................................................................................................... 3
Echinoids ......................................................................................................................................................... 6
Ophiuroids....................................................................................................................................................... 8
Asteroids ....................................................................................................................................................... 10
Holothurians.................................................................................................................................................. 16
Appendix 1 ................................................................................. 47
List of codes based on the classification of Parker (1982)
Appendix 2 ................................................................................48
List of international specialists consulted
Appendix 3 ................................................................................52
Identification, geographic and bathymetric distribution of echinoderms collected
in the Pacific Ocean
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IOC TECHNICAL SERIES NO. 69, VOL.2
The Echinoderms
of the Clarion-Clipperton
fracture zone
T
he method of identification used in this photographic atlas, based on descriptions of organisms
observed in situ, has already been used in several publications, notably those by Macurda & Meyer (1976),
Roux (1980), Pawson (1983), Ohta (1985), Pawson & Foell
(1986), Fujita et al. (1987), Pawson (1988), Miller & Pawson
(1990) and Bluhm & Gebruk (1999). Most of the species
of this zone are probably new to science since this part
of the Pacific Ocean has scarcely been sampled. Also the
in situ appearance of the megafauna differs considerably
from that of preserved specimens, and especially so for
the echinoderms, which often loose their superfical external gelatinous layer and delicate appendages at the
time of collection (Bluhm & Gebruk, 1999). Photographic
and video material are also the only means that allow the
collection of valuable data on the behaviour and environment of each taxon identified in the midst of the faunal
assemblages which compose the benthic and suprabenthic megafauna (Owen et al., 1967; Rowe, 1971; Grassle et
al., 1975; Lemche et al., 1976; Cohen & Pawson, 1977; Patil
et al., 1980; Wigley & Theroux, 1981; Mauviel, 1982; Ohta,
1985; Laubier et al., 1985; Foell, 1988; Pawson, 1988a, b;
Tilot et al., 1988; Foell & Pawson, 1989; Kaufmann et al.,
1989; Wheatcroft et al., 1989; Bluhm, 1991; Thiel et al.,
1991; Sharma & Rao, 1991; Christiansen & Thiel, 1992;
Smith et al., 1992; Christiansen, 1993; Bluhm, 1994; Bluhm & Thiel, 1996; Lauerman et al., 1996; Radziejewska,
1997; Piepenburg & Schmid, 1997; Hughes & Atkinson,
1997; Fukushima & Imajima, 1997; Kaufmann & Smith,
1997; Kotlinski & Tkatchenko, 1997; Matsui et al., 1997;
Tkatchenko & Radziejewska, 1998; Nybakken et al., 1999;
Radziejewska & Kotlinski, 2002).
Often the study of the megafauna is limited to particular
zoological groups such as echinoderms (Haedrich et al.,
1980; Sibuet & Lawrence, 1981; Rice et al., 1982; Pawson,
1983; Briggs et al., 1996; Copley et al., 1996; Piepenburg
et al., 1996; Bluhm & Gebruk, 1999) and sometimes limited to holothurians (Matsui et al., 1997). Echinoderms and
especially holothurians are also known to be good indicator species in impact studies on the exploitation of polymetallic nodules deposits (Thiel et al., 1992; Bluhm et al.,
1995; Bluhm & Gebruk, 1999) or in studies of the variation
in particle flux of organic carbon (Smith et al., 1997).
The echinoderms of the Clarion-Clipperton fracture zone
are presented here in the form of a photographic atlas
accompanied by a commentary for each taxon. The gear
that provided the photography and video of the seabed
involved towed and free-moving vehicles:
The towed vehicles, generally multi-instrumented, are:
• the “Remorquage Abyssal d’Instrumentation pour
l’Exploration” or “R.A.I.E.” developed for Ifremer
• an epibenthic camera sled or “troika” perfected by
Commander Cousteau
• The “Deep Tow Instrumentation System” or “DeepTow” developed for the Scripps Institute of oceanography (USA)
The remote devices, all developed by Ifremer, are:
• the automatic sampler coupled to a camera or
“E.D.1”
• the unmanned free-moving submersible “Epaulard”
• the manned deep-diving submersible “the Nautile”
In the photographic atlas, each commentary is presented in the following order:
• a morphological description,
• a description of certain aspects of behaviour (such
as methods of locomotion, visible tracks, swimming
movements, trophic behaviour ...),
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• edaphic preferences (the occurrence of a taxon on a
particular substratum or nodule-facies),
• hydrological preferences (such as orientation with
respect to currents)
• hypotheses of identification made in agreement with
international specialists for each group,
• information on morphology extracted from the literature,
• bathymetric and geographic distributions for each
taxon in relation to the proposed identification(s),
• comparative observations in relation to taxa identified from other sites in the Clarion- Clipperton fracture zone, such as the DOMES sites, and more precisely
the Echo 1 site located near DOMES C (14°40’N125°25’W). These sites were investigated by the National Oceanic and Atmospheric Administration (NOAA)
and the Scripps Institution of Oceanography. Similarities are also noted with the taxa identified from a
station further north-east (34°50’N, 123°00’W), near
the central part of the Californian coast at 4 100 m
depth (Lauerman et al., 1996; Kaufmann & Smith,
1997). Other sites of comparison outside the ClarionClipperton fracture zone are in a polymetallic nodules
area in the abyssal Peruvian Basin in the south-east
Pacific Ocean (Bluhm, 1994; Bluhm & Gebruk, 1999)
Crinoids
CRI 1
This species often has five arms (figure 1) or a multiple
of five arms, on which pinnules can sometimes be distinguished. The arms measure between 10 and 20 cm.
The longest arm in Figure 1 is 20 cm and the crown of
arms is spreading, as a “feeding net”, to trap food particles efficiently. This species has also been observed
moving around by the alternating motion of its arms. In
this figure, its star-shaped tracks can be seen situated somewhat below the animal, as though produced during
its displacement.
This species was observed on the facies of polymetallic
nodules as well as on purely sedimentary substratum,
facies O, as in Figure 1.
and in a totally different abyssal environment at similar depth, in the Bay of Biscay in the north-east Atlantic Ocean (Sibuet et al., 1980).
• Some remarks on the preceding ideas may conclude
each commentary.
The list of codes used in the identifications, following
the classification of Parker (1982), is presented as Appendix 1. The international specialists on whose advice
the hypotheses of identification in this study were compiled are listed in Appendix 2. The accepted hypotheses
of identification are presented and commented upon in
the photographic atlas. Based on these, Appendix 3 presents, in the form of a table, a synthesis of information
from both specialists and the literature on the echinoderms sampled in the Clarion-Clipperton fracture zone
or in the eastern Pacific Ocean region. Also for each species identified are presented the author and date of identification, the bathymetric and geographic distributions
and the substrata conditions in which it was collected.
The last column displays the codes for the taxa and the
name of the international specialists who participated in
their identification.
this free-living crinoid belongs to the family Antedonidae, Order Comatulida, subclass Articulata, class Crinoidea. The hypothesis of taxonomic identification for
this comatulid is Fariometra parvula (Hartlaub, 1895),
according to the specimens identified by Hartlaub (1895)
and Clark (1923, 1967) between latitudes 32°N and 2°N in
the Eastern Pacific Ocean, at depths between 589 m and
1 969 m and on substrata of green mud and globigerina
ooze.
The family Antedonidae is represented in the Atlantic Ocean (Sibuet et al., 1980) and the Pacific Oceans
between 1 200 m and 5 000 m (Carpenter, 1888). This
species was listed by Lauerman et al. (1996) from a site in
the north-eastern Pacific Ocean close to the central part
of the Californian coast at 4 100 m depth and it has also
been observed in the Peruvian Basin (Bluhm, 1994).
In reference to the literature (Carpenter, 1888; Hartlaub,
1895; Clark, 1923, 1967) and in the opinion of specialists,
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CRI 2
CRI 3
These crinoids are yellow-orange in situ as in photographs taken by the “Nautile”. Their open crown, formed of
10 pinnate arms, is 20 cm diameter and may be parallel
to the substratum or flattened by the current. As rheophiles, they feed by the passive filtering of suspended
particles (Roux, 1980). Their long peduncle, to which
small actiniae or cirripeds are often attached, is anchored either in sediment or more often on a hard substratum such as rock outcrops or nodules (figure 2).
This small sedentary crinoid has a peduncle 10 cm high
and a crown of five arms fringed with long pinnules (figures 3, 4). Each arm measures a maximum of 5 or 6 cm.
According to the literature (Carpenter, 1884; Clark, 1908;
Roux, 1980; Luke, 1982), these fixed crinoids could be
Bathycrinus equatorialis (Clark, 1908), a representative
of the family Bathycrinidae, Order Bourgueticrinida.
Pawson & Foell (1983) suggested Rhizocrinus sp. or
Bathycrinus sp. for the taxa they photographed at the
DOMES C site in the Clarion-Clipperton fracture zone.
This taxon was also listed by Lauerman et al. (1996) at
a site in the north-east Pacific close to the central part
of the Californian coast at 4 100 m depth.
In view of the difficulties of distinguishing differences
in photographs, specialists have also suggested that
this crinoid could perhaps be Bathycrinus affinis (Roux,
1980), Bathycrinus aff. australis (Roux, 1980), Rhizocrinus sp. (Carpenter, 1884) or Monachocrinus sp. (Clark,
1917). The latter has been reported at 32°N, on the Patton Escarpment, between 3 600 m and 3 676 m (Luke,
1982). These crinoids occur in the Atlantic, Indian and
Pacific Oceans at a great variety of depths (Carpenter,
1884).
Similar sedentary crinoids, but very dark in colour and
larger, were identified as Bathycrinus sp. in photographs from the Peruvian Basin (Bluhm, 1994). The Bathycrinids (Bathycrinus gracilis, Monachocrinus recuperatus, Zeuctocrinus gisleni) identified by Roux (1985)
in the north-east Atlantic Ocean between 2 000 m and
5 000 m are represented in photographs from the Biogas cruise (Sibuet et al., 1980) and the Cymor cruise
between 49°N and 30°N on the continental margin of
the Bay of Biscay.
According to specialists and information from the literature (Carpenter, 1884; Clark, 1907; Roux, 1980), the proposed identification is Ptilocrinus sp. (Clark, 1907), family
Hyocrinidae, Order Millericrinida. Macurda & Meyer (1976)
described a dense pinnulation specific to Ptilocrinus sp..
Some specialists are undecided between Hyocrinus sp.
and Ptilocrinus sp. (Roux, 1980).
Based on photographs from the DOMES C site, Pawson &
Foell (1983) proposed the identification Hyocrinus bethellianus (Wyville Thomson, 1876), which has a wide distribution from the equatorial Atlantic Ocean to the Crozet Islands in the Mid-Indian Ocean between 2 800 m and 3 300
m (Carpenter, 1884). This yellow hyocrinid has a smooth
peduncle composed of ossicles of calcium carbonate
which are shaped like dumbbells. The pinnules decrease in
size distally along the arms. This hyocrinid was not found
in the photographic analyses of Bluhm (1994) for the Peruvian Basin.
The crinoids identified by Roux (1980) as being Hyocrinus
sp. were collected from a rocky substratum on the East
Pacific ridge to the west of theTres Marias Islands between
2 646 m and 2 665 m depth. A new species belonging to
the family Hyocrinidae, Laubiericrinus pentagonalis, was
described by Roux (2004) from specimens collected by
submersible at 2 765 m depth in the south-west Pacific
Ocean.
The Hyocrinidae Anachalypsicrinus nefertiti and Hyocrinus
grimaldii were identified in photographs from the Cymor
cruise exploring the Armorican margin at about 2 500 m
depth and the Biogas cruise in the Bay of Biscay at about
3 000 m depth (Sibuet et al., 1980; Roux, 1985).
The collected crinoids, which on the advice of the specialists appear similar to the taxa identified in the ClarionClipperton fracture zone, are presented in appendix 3 with
their geographic and bathymetric distribution.
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Fig. 1. Fig. 1. Photograph taken by the ‘Epaulard’ at the NIXO
45 site: nodule-facies O, crinoid Fariometra parvula (CRI 1)
© Ifremer
Fig. 4. Photograph taken by ‘Deep Tow’ at the DOMES C site:
mixed nodule-facies, crinoid Hyocrinus bethellianus (CRI 3)
© Ifremer
Fig. 2. Photograph taken by the “E.D.1”
device during the Copano cruise: nodule-facies C+, crinoid Bathycrinus sp. (CRI 2) and
echinoid Pleisiodiadema globulosum (OUR 1)
© Ifremer
Fig. 3. Photograph
taken by the ‘Epaulard’ at
the NIXO 45 site: crinoid
Hyocrinus bethellianus
on a cliff (CRI 3)
© Ifremer
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Echinoids
OUR 2
OUR 1
One sees here a very sinuous track which could have
been created by an animal living in the sediment. Similar tracks from the Peruvian Basin have been identified
as having been made by irregular echinids (figure 6).
This echinid with long slightly recurved radioles about
10 to 15 cm long has a body 2 to 5 cm diameter (figure
5). In the images taken by the ‘Nautile’ it appears violet,
often with a paler region on the aboral face (lightly covered with sediment, reminiscent of the camouflage commonly found in littoral species). OUR 1 was observed on
substrata with sediments and nodules as well as on the
ocean floor in the Peruvian Basin.
Based on the literature (Agassiz, 1898, 1904, 1908; Mortensen, 1903, 1940, 1951; Downey, 1968) and the advice
of specialists, the hypothesis of identification for this
taxon is Pleisiodiadema globulosum (Agassiz, 1898),
family Aspidodiadematidae, Order Diadematoida. It is
a regular sea urchin which shares the abyssal environment of the Pacific and Atlantic Oceans with another genus, Aspidodiadema (Agassiz, 1898).
In the eastern Pacific Ocean, Pleisiodiadema globulosum
has been collected between 8°N and 10°S, between Malpelo Island and Valparaiso from depths between 2830
m and 3900 m. This echinid was identified by Pawson &
Foell (1983) from the Domes C site in the Clarion-Clipperton fracture zone, and in the inventory of the megafauna
of the Peruvian Basin, it was also present, along with
Plesiodiana sp. and at least three other undetermined
echinids (Bluhm, 1994).
The family Aspidodiadematidae is characterised by a
very fragile, globular test and long brittle spines (often
in photographs, the echinids have radioles that are shorter in certain places, probably having been broken and
in the process of regeneration). The radioles of the adoral side in contact with the benthos are terminated by
a suction pad, in the form of a club that has a locomotary function, while those on the aboral surface end in a
point and are specialised for respiratory and tactile roles
(Cuénot, 1948). The presence of a weak shadow next to
the animal suggests that it is raised up on its adoral radioles.
This irregular echinid has a fragile oral skeleton with a
depressed region on the aboral surface encircled by a
well-defined narrow brown band. The podia with terminal dics are localised in this depressed region and their
function is similar to that of the sweeping podia of numerous burrowing spatangoids, which construct a chimney that communicates between the open water and
the burrow (Smith, 1980). This irregular echinid has been
identified as Aceste ovata (Agassiz & Clark, 1907), family
Aeropsidae, suborder Aphisternata, Order Spatangoida.
Since this sea urchin has an endobenthic way of life,
its white skeleton can only be partially seen, lying on
the sediment when it is dead, and it then resembles an
ovoid sponge.
Pawson & Foell (1983) identified this echinid in images
and from specimens collected at the DOMES C site in
the Clarion-Clipperton region. It has also been observed
near Hawaii, off Japan and in the Indian Ocean at depths
between 450 m and 4 800 m (Koehler, 1914), as well as
amongst the megafauna of the Peruvian Basin (Bluhm,
1994).
The spatangoids are heart-shaped sea urchins that feed
on mud or are microphages. Their spines are very small
and numerous. The cosmopolitain family Aeropsidae is
exclusively abyssal and bathyal. It contains only two genera, Aeropsis and Aceste, collected in particular in the
Bay of Biscay (Cuénot, 1948; David & Sibuet, 1985).
Lauerman et al. (1996) and Kaufman & Smith (1997)
idenitifed an echinoid Echinocrepis sp. at 4 100 m at a
station further north and close to the central part of the
Californian coast.
The other regular and irregular echinids that can be observed and identified in the Clarion-Clipperton region
are listed in appendix 3 with their geographic and bathymetric distribution.
It is possible to observe in figure 5 the bioturbulent activity of these oral radioles on the substratum, uncovering
a paler sediment and providing evidence of non-selective feeding behaviour in OUR 1, by ingesting sediment
as well as detritus.
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Fig. 5. Photograph taken by the “E.D.1” device during the Copano cruise: nodule-facies C+
sparse, echinoid Pleisiodiadema globulosum (OUR 1) © Ifremer
Fig. 6. Photograph taken by the ‘Epaulard’ at the NIXO 45 site: nodule-facies C, tacks of irregular echinoid Aceste ovata
(OUR 2) © Ifremer
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Ophiuroids
OPH 1
This species is 6-14 cm diameter with a very white, flat
central disc and five spindly arms, which are 4-5 times
as long as the diameter of the central disc (figures 7 and
9). These taxa are found on all forms of substratum from
rock outcrops to sediments with or without nodules.
In agreement with specialists and after referring to the literature (Lyman, 1882; Lütken & Mortensen, 1899; Clark,
1915; Koehler, 1922; Downey, 1969), the proposed identification is Ophiomusium armatum (Koehler, 1922), family Ophiuridae, suborder Chilophiurina, Order Ophiurida, subclass Ophiuroidea, class Stelleroidea. Members
of the family Ophiuridae are relatively large with long,
slender arms, and this family presents the characteristics of the true ophiurids, in which the arms are never
branched and not readily moved except in a horizontal
plane. The discriminant characters for this family are
only visible by microscope. The ophiurids are one of the
most ancient forms, dating from the Ordovician. They
are cosmopolitan in abyssal environments with more
than 30 species in the genus Ophiomusium. According
to the specialists, OPH 1 could equally belong to the genus Ophiomusium, family Ophiuridae, or to the genus
Amphiophiura, family Amphiuridae, or to either Ophiura
or Stegophiura, although Ophiomusium has the longest
legs. Superficially, all these ophiurids closely resemble
one another.
Several undetermined forms of ophiuroids have been
reported from the Clarion-Clipperton fracture zone by
Pawson & Foell (1983) and from the Peruvian Basin by
Bluhm (1994). Lauerman et al. (1996) and Kaufman &
Smith (1997) identified ophiuroids at a site in the northeast Pacific Ocean close to the central part of the Californian coast at 4100 m depth as Ophiura bathybia, Amphilepsis patens and Amphiura carchara on sediments, and
Ophiacantha spp. on rocky substrata.
ophiuroids could be suspension feeders, based on observations of Antarctic specimens that produce a thread
of mucus which is held between the spines of the arms
in order to trap the small nekton, transported towards
the mouth by movements of the flagella. Another way
of being a suspension feeder, as in the case described
by Fujita & Ohta (1988) for ophiuroids associated with
gorgonians, involves capturing particles in suspension,
transported by currents. In the Clarion-Clipperton study
zone an association was frequently seen between ophiurids and sponges of the genus Hyalonema, which are attached by long peduncles. This would confirm the hypothesis of Fujita & Ohta (1988) of suspension feeding by
the ophiuroids of the Clarion-Clipperton fracture zone.
OPH 1’
According to the specialists, this ophiuroid is a dark version of the same genus as the preceding taxon, Ophiomusium armatum (Koehler, 1922). It appears smaller
and more slender, and is difficult to identify from photographs (figure 8).
Another ophiure, Amphiophiura convexa (Lyman,
1878), family Ophiuridae, suborder Chilophiurina, Order
Ophiurida, has been identified from the Clarion-Clipperton Zone (Pawson & Foell, 1983). It is characterised by
short arms that taper rapidly and bear three small spines on each side of the joints. The aboral surface of the
central disc comprises large plates surrounding a single broad central plate. This generally small brittle star
is difficult to detect in photographs and in addition, it
generally lives buried close to the water-sediment interface. This taxon has been collected in many locations in
the northern Atlantic Ocean, in the western part of the
Indian Ocean and in the Arabian Sea at depths between
1 997 m and 6 810 m (Lyman, 1882; Bortsch, 1983; Vadan
& Guille, 1984).
The collected ophiurids, which according to specialists
appear similar to the taxa identified in the Clarion-Clipperton fracture zone, are given in Appendix 3 with their
geographic and bathymetric distribution.
Various ideas on the feeding behaviour of these abyssal
brittle stars have been proposed in the literature. According to David & Sibuet (1985), they selectively ingest
detritus and sediment particles, whereas Litvinova & Sokolova (1971) and Pearson & Gage (1984) consider them
to be non-selective euryphagous (omnivores), which are
motile and opportunist, feeding on all prey or detritus
found in their path. On the other hand, Sokolova (1972)
says that abyssal cosmopolitan brittle stars are essentially carnivores, and in particular the semi-microphagous species. However, Fell (1961) suggests that some
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Fig. 7. Photograph taken by ‘Deep Tow’ at the Echo I site: nodule-facies C, ophiuroid Ophiomusium armatum (OPH 1) © Ifremer
Fig. 8. Photograph of a sample from the Copano
cruise, dark ophiuroid Amphiophiura convexa
(OPH 1’) © Ifremer
Fig. 9. Photograph taken by the ‘Nautile’ during the NIXONAUT cruise: nod-
ule-facies C sparse, ophiuroid Ophiomusium armatum (OPH 1)
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Asteroids
AST 1
This asteroid is about 12 cm diameter, with thick, almost
cylindrical arms that lack marginal spines, a slightly
prominent disc and a somewhat rough appearance
(figure 10). In some photographs, the central disc can
sometimes be very swollen due to ingested sediment,
indicating a detritvorous feeding habit. These asteroids
are most often observed on substratum with a dense covering of nodules.
In agreement with specialists and after referring to the
literature (Fisher, 1905, 1911, 1928, 1930; Ludwig, 1905;
Clark, 1913, 1923; Madsen, 1951; Downey, 1970; Luke,
1982), the proposed identification is a representative
of the family Zoroasteridae, in the Order Forcipulatida,
either Zoroaster hirsutus (Ludwig, 1905) or Zoroaster
ophiurus (Fisher, 1905). The latter has been collected in
the south-eastern Pacific Ocean (Fisher, 1905) but has
not been recorded in the photographs from the Peruvian
Basin (Bluhm, 1994).
AST 2
This asteroid is about 15 cm diameter and is generally
buried, leaving only its central disc of about 5 cm diameter prominent (figures 11 and 12). Pentahedral tracks on
the circular mounds in which they are buried can probably be attributed to them. Similar asteroids have been
observed in the Bay of Biscay (Sibuet et al., 1980).
Based on the literature (Sladen, 1883; Ludwig, 1905,
1907; Fisher, 1955; Madsen, 1961; Luke, 1982; Belyaev,
1985) and in agreement with specialists, the proposed
identification for this taxon is a member of the family
Porcellanasteridae, Order Paxillosida. Some specialists
have suggested Hyphalaster inermis (Sladen, 1883),
which has been reported in the Clarion-Clipperton region
(Pawson & Foell, 1983), as have the following Porcellanasterids: Eremicaster pacificus (Ludwig, 1905), Eremicaster crassus gracilis (Sladen, 1883) and Thoracaster
cylindratus (Sladen, 1983) (cf. Appendix 3). This asteroid
has not been recorded in photographs from the Peruvian Basin (Bluhm, 1994).
Although they do not appear in the photographs, the
distinctive characteristics of Hyphalaster inermis are
the spines at the rounded extremity of the arms and
the aboral marginal plates which join the arms midway
(Madsen, 1961).
Because of the burrowing nature of this animal, which
is rarely visible at the sediment surface, the distinctive
characters of Thoracaster cylindratus (Sladen, 1883), i.e.
the two single marginal plates on the dorsal and ventral
sides at the centre of each inter-radius, cannot be seen
either. In fact one part of the aboral surface is kept out of
the sediment due to the presence of paxillae (small erect
columns on the plates of the test that bear spreading
spines at their free end), which on contact draw together
to form an impenetrable roof (Fell, 1982).
AST 3
This asteroid has a fairly well developed disc and rigid,
tapering arms with pointed ends. Its diameter is about
15 cm and it is seen especially on sedimentary substrata.
In accordance with the specialists and by reference to
the literature (Ludwig, 1905; Fisher, 1910, 1911; Clark,
1913, 1920, 1923; Clark, 1981; Luke, 1982), the proposed
identification of this asteroid is Pectinaster sp., a representative of the family Benthopectinidae, Order Paxillosida. Sometimes aggregates can be distinguished along
its arms, which could be attached to the well-developed
spines found on the aboral marginals (Fell, 1982).
This taxon resembles Pectinaster agassizii (Ludwig,
1905), sampled notably in the north-east Pacific Ocean
at medium depths (Clark, 1981), as well as another Benthopectinid, Benthopecten acanthonotus (Fisher, 1905),
which has been collected in the tropical east Pacific
Ocean between 1 157 m and 2 761 m (cf. appendix 3).
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Fig. 10. Photograph taken by
the ‘Epaulard’ at the NIXO 45 site: nodule-facies C, asteroid Zoroaster hirsutus
(AST 1) © Ifremer
Fig. 11. Photograph taken by the ‘Epaulard’ at the
NIXO 45 site: rock outcrop and nodule-facies O, asteroid Hyphalaster inermis (AST 2)
© Ifremer
Fig.12. Photograph taken by the ‘Epaulard’
at the NIXO 45 site: nodule-facies O, asteroids
Hyphalaster inermis (AST 2) and Hymenaster
violaceus (AST 5)
© Ifremer
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AST 4
AST 5
This species is about 5 cm diameter with a pentagonal
body. It was mostly observed on ocean floor with a dense covering of polymetallic nodules. It is present in some
photographs from the Peruvian Basin (Bluhm, 1994).
In figures 12, 13 and 14, an asteroid 5 to 10 cm diameter with a fleshy membrane can be distinguished. This
species is sometimes violet in photographs taken by the
“Nautile”.
According to the literature (Ludwig, 1905; Clark, 1920;
Madsen, 1951) and in agreement with the specialists, the
pentahedral shape of this asteroid and the translucent
membrane on the aboral surface are typical of representatives of the family Pterasteridae, in the Order Spinulosida. This level of identification must suffice as the
elements for greater precision are lacking. In some photographs, long, fine, prickly adambulacra can be seen,
perpendicular to the arms and enclosed in the membrane. In typical members of this family, with the exception
of Hymenaster, the prickles on the adambulacra subtend
two lateral palmate membranes (Fell, 1982).
According to the literature (Ludwig, 1905; Clark, 1920;
Madsen, 1956) and in agreement with the specialists,
the proposed identification for this taxon is Hymenaster
violaceus (Ludwig, 1905). The distinctive features of the
genus Hymenaster given by Sladen (1889) and Mortensen (1927) are a supradorsal membrane composed of
numerous muscular fibres, well developed actinal prickles covered by a membrane, and adambulacral prickles
that are free and not palmate.
This species has been reported from the DOMES C site
(Pawson & Foell, 1983) and in the Peruvian Basin (Bluhm, 1994), as well as on rocky facies explored during the
Cymor cruise (North Atlantic).
This asteroid might be expected to have omnivorous
feeding habits, both detritivorous and carnivorous,
based on the examination of the gastric contents of Hymenaster sp. and Hymenaster quadrispinosus (Fisher,
1905) collected at 1 600 m and 2 926 m (Carey, 1972).
However, it may instead be a predominantly detritivorous opportunist, as many abyssal organisms are (Sokolova, 1959). This asteroid colonizes sediments as well as
rocky substrata (Fisher, 1911; Clark, 1920; Madsen, 1951)
and in photographs of the study zone, it was only found
on substrata with polymetallic nodules.
Hymenaster quadrispinosus (Fisher, 1905), Hymenaster gracilis (Ludwig, 1905) and Hymenaster violaceus
(Ludwig, 1905) have been collected at depths between
1 935 m and 3 436 m in the north, central and south-eastern Pacific Ocean (Appendix 3).
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Fig. 13. Photographie prise par le «Deep
Tow» sur le site d’Echo I : Faciès-nodules B,
astéride Hymenaster violaceus (AST 5)
Fig.14.
Photograph taken by
the ‘Nautile’ during
the NIXONAUT
cruise: nodule-facies
C, asteroid Hymenaster violaceus
(AST 5)
© Ifremer
Fig. 15. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
nodule-facies C, brisingid asteroid
with 6 arms captured by the ‘Nautile’,
Freyella benthophila (AST 6)
© Ifremer
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AST 6
AST 7
These asteroids have six arms, each measuring 5 to 20
cm in length (figures 15, 16, 17). The arms are very thin
and clearly distinct from a small central disc, which gives this taxon the appearance of a brittle star. White or
yellow-orange in colour, Freyella spp. are generally quite
abundant and attached to nodules.
Figure 18 represents a Brisingidae with seven arms,
each measuring about 20 cm. Freyella species with
seven arms have also observed in the Peruvian Basin
(Bluhm, 1994).
Similar Brisingidae have been reported in the ClarionClipperton fracture zone ((Pawson & Foell, 1983) and in
the Peruvian Basin (Bluhm, 1994).
Based on information from the literature (Sladen, 1889;
Clark, 1920; Fisher 1928, 1930; Madsen, 1951, 1956) and
the advice of specialists, the proposed identification for
this asteroid is a member of the genus Freyella, family
Brisingidae, Order Euclasterida (Downey, 1986). Freyella
is cosmopolitan in abyssal environments and can have
six or more arms. Some specialists have suggested this
taxon might be Freyella benthophila (Sladen, 1889; Clark,
1920a) whose distribution extends across the eastern Pacific Ocean to the Indian Ocean in abyssal environments
(Madsen, 1951, 1956; Fisher, 1928, 1930). Cherbonnier &
Sibuet (1972) have also collected and identified Freyella
benthophila in the Atlantic (45°13’N-5°30’W) at 4 700 m
depth.
The majority of Freyella observed in the study zone had six
arms but other Brisingidae had five, seven or eight arms
(Freyella octoradiata, Clark, 1920), nine arms (Freyella heroina, Sladen, 1889) or 10 arms. The problem of number
of arms as a means of determination in Freyella is tricky
when one considers the frequency of autotomies and
the regenerative power of a variable number of brachial
buds, as in all the non-pentagonal asteroids which do not
have arms largely united to the disc (Cuénot, 1948). Some
specialists suggested that AST 6 could be a Freyastera
(Downey, 1986). A Freyella with five arms has been observed in a single photograph from the Clarion-Clipperton
fracture zone and could be an accidental case.
Some behavioural observations could be made on the
feeding posture of this brisingid asteroid. Freyella generally has its arms slightly upturned in order to present the
natural filter of their adherent podia to the current, thus
trapping particles in suspension. This posture is similar to
that of the comatulids. The position of the arms also indicates the direction of the current.
The Freyella sp. in figure 18 has a less typical posture,
with its arms stretched out and their extremities sometimes raised. The arms are fringed with long thin podia
that comb the water-sediment interface. This difference
in posture could indicate a new species of Freyella (Pawson, 1982) or another, more detrititus feeding behaviour.
Note also that this posture demonstrates maximum efficiency in the pursuit of food by dividing up equally the
area to be searched.
According to Downey (1986), the Brisingidae can be separated from the Freyellidae by their feeding behaviour,
the Brinsingidae having their arms upturned in the current while the Freyellidae have theirs extended over the
substratum. However, this statement does not seem justified as photographs from the Clarion-Clipperton fracture zone show that both taxa can have their arms in
either position.
AST 8
Figure 19 shows a Brisingidae of the same diameter as
AST 7 with 10 arms. As in all the other Freyella species
observed in the Clarion-Clipperton fracture zone, they
are found preferentially on nodules. Freyella with 11
arms were seen in photographs from the Bay of Biscay
(Sibuet et al., 1980).
By common agreement of the specialists and with reference to the literature (Sladen, 1889; Clark, 1920; Fisher
1928, 1930; Madsen, 1951, 1956), the proposed identification is Freyella sp. because neither Freyella brevispina
(Clark, 1920) nor Freyella insignis (Ludwig, 1910), collected in the eastrn Pacific Ocean, ever have 10 arms, but 11
and 13 respectively (A. Clark, pers. comm.).
The collected asteroids, which in the opinion of the specialists appear similar to the taxa identified in the Clarion-Clipperton fracture zone, are listed in Appendix 3
with their geographic and bathymetric distribution.
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Fig.16. Photograph taken by
the ‘Epaulard’ at the NIXO 45 site: mixed
nodule-facies B and rock outcrop, brisingid
asteroid with 6 arms, Freyella benthophila
(AST 6) © Ifremer
Fig. 17. Photograph taken
by the ‘Epaulard’ at the NIXO
45 site: nodule-facies O,
brisingid asteroid with 6 arms,
Freyella benthophila (AST 6)
© Ifremer
Fig. 18. Photograph taken by
‘Deep Tow’ at the Echo I site: nodule-facies C,
asteroid with 7 arms Freyella sp. (AST 7)
© Ifremer
Fig. 19. Photograph taken by the ‘Epaulard’ at the NIXO 43 site:
mixed nodule-facies (A and C), asteroid with 10 arms Freyella sp.
(AST 8) © Ifremer
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Holothurians
HOL 1
This whitish holothurian measures 10 to 20 cm in length
by 2 to 3 cm wide (figures 20, 21, 22). Its narrow body
with attenuated extremities appears cylindrical with one
ventral, plantar surface. Its rigid appearance suggests a
thick cuticle, which is scarcely bristly with at least two
rows of short, rudimentary dorsal papillae, sometimes
irregularly arranged. In the study zone, this taxon was
always relatively abundant, never seen in the water column and apparently indifferent to the type of substratum and sediments on which it feeds, and whether or
not they are covered with nodules.
Similar Synallactes species were recorded by Pawson
& Foell (1983) and Lauerman et al. (1996), as well as by
Kaufman & Smith (1997) at a site in the north-eastern
Pacific Ocean near the central section of the Californian
coast at 4 100 m depth. This orange to brown taxon has
also been found at great depths in the Peruvian Basin
where they were identified as Synallactes sp. type 2
(Bluhm, 1994; Bluhm & Gebruk, 1999).
On the advice of specialists (Pawson, Sibuet, pers.
comm.) and after referring to the literature (Ludwig,
1894; Clark, 1920; Madsen, 1953; Hansen, 1975), the identification proposed is a taxon belonging to the family
Synallactidae, Order Aspidochirotida. The quality of the
photographs did not allow further analysis and the recognition of Synallactes aenigma (Ludwig, 1894), which
has been collected in the north-central Pacific Ocean (appendix 3). Bluhm & Gebruk (1999) suggested Synallactes
reticulatus (Sluiter, 1901) as a second possible name for
this taxon observed in the Peruvian Basin.
HOL 2
This whitish holothurian has a narrow, cylindrical body
(figures 23, 24). Its dorsal surface is covered by at least
two rows of numerous slender papillae of variable size
but not however exceeding the width of the body. This
species measures 12 to 20 cm long and 2 to 4 cm wide.
Just as with the preceding taxon, this holothurian is
found on all types of substratum, whether covered or
not by nodules. It is not known to be mesopelagic (cf.
appendix 3).
This holothurian is fairly cosmopolitan having been reported at depths between 2 700 m and 4 300 m in the
Indian, Pacific and Atlantic Oceans on a great variety of
substrata. Pawson & Foell (1983) identified it at the DOMES C site, and Kaufman & Smith (1997) also identified
it. It has also been found with a pink tinge in the abyssal
depths of the Peruvian Basin (Bluhm & Gebruk, 1999),
who referred to it as Synallactes sp. type 1. This taxon
has also been reported in the Atlantic Ocean, notably in
the Bay of Biscay (Sibuet et al., 1980).
The identification proposed in accordance with the literature (Koehler & Vaney, 1905; Clark, 1920; Madsen, 1953;
Hansen, 1975) and specialists (Pawson, Massin, pers.
comm.) is Synallactes profundi (Koehler & Vaney, 1905)
in the family Synallactidae and Order Aspidochirotida.
This taxon has been collected in the Indian, central and
south Pacific and north Atlantic Oceans (appendix 3).
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Fig. 20. Photograph taken by the ‘Epaulard’ at the NIXO 45 site:
nodule-facies C, holothurian Synallactes aenigma (HOL 1)
© Ifremer
Fig. 21. Photograph taken by a “Troika” during the Copano 1 cruise:
nodule-facies C, holothurian Synallactes aenigma (HOL 1)
© Ifremer
Fig. 22. Photograph taken by the ‘R.A.I.E.’ at the NIXO 45 site:
nodule-facies C, holothurian Synallactes aenigma (HOL 1)
© Ifremer
Fig. 23. Photograph
taken by the ‘Epaulard’ at the NIXO 45
site: nodule-facies C,
holothurian Synallactes
profundi (HOL 2)
© Ifremer
Fig. 24. Photograph taken by the ‘Epaulard’ at
the NIXO 45 site: mixed nodule-facies (C and B),
holothurian Synallactes profundi (HOL 2)
© Ifremer
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HOL 3
HOL 4
This white-coloured taxon can reach 25 cm long and 6
cm wide (figures 25, 26). It is remarkable in its elongated semi-cylindrical form with rounded extremities. The
dorsal papillae are relatively long, thin and aligned in
one row per radius, at the interior of which they are uniformly distributed. These papillae are rigid at the base
and more flexible at the ends. This holothurian is is not
mesopelagic, it is generally found on substrata with a
dense covering of nodules and it. It leaves characteristic tracks on the sediment which consist of a double
row of small depressions that infer the existence of a
double row of ventral podia and the absence of medioventral podia.
This holothurian measures a maximum of 20 cm long
and 4 cm wide, and is white with a rigid, cylindrical
body (figures 27, 28). It closely resembles HOL 3 but is
more thickset. Its dorsal surface has more numerous,
long, delicate, rigid papillae distributed in several rows,
with 4 to 35 pairs per dorsal row and 4 to 17 pairs per
lateral row. The trail it creates makes a double row of
small holes, indicating the absence of medio-ventral
podia. This taxon has been observed on all types of
substrata. It appears to ingest detritus and sediments
in a non-selective manner, and is not known to be able
to move in the water column.
Pawson & Foell (1983) identified this holothurian from
the DOMES C site, however, it does not appear in the
identifications of Lauerman et al. (1996) nor of Kaufmann & Smith (1997). It was not reported from photographs of the depths of the Peruvian Basin (Bluhm,
1994). It is poorly represented in the Bay of Biscay
(Sibuet et al., 1980).
By reference to the literature (Théel, 1879; Ludwig, 1893,
1894; Clark, 1920; Madsen, 1953; Hansen, 1975) and in
agreement with the specialists, we suggest that this
taxon belongs to the genus Orphnurgus (Théel, 1879) in
the family Deimatidae and Order Elasipodida. This holothurian demonstrates the visible characters of a member
of the family Deimatidae by the presence of numerous
long papillae, the mouth situated in the anterior part of
the ventral sole and podia only on the external radius
of the trivium. The genus Orphnurgus, previously known
as Scotodeima (Ludwig, 1894), is characterised by the
presence of 15 to 20 non-retractable tentacles and by the
absence of peribuccal papillae.
Pawson & Foell (1983) identified this holothurian from
the DOMES C site, and Lauerman et al. (1996) and Kaufman & Smith (1997) have also identified it from 4100 m
at a site further north. It has been recorded from the Peruvian Basin (Bluhm, 1994; Bluhm & Gebruk, 1999) but
appears somewhat different in their photographs, being
narrower with two longer papillae on the anterior part
of the body and two other perpendicular papillae on the
dorsal posterior part. This taxon has been found also in
the Bay of Biscay (Sibuet et al., 1980).
The proposed identification, after consulting the literature (Théel, 1879; Ludwig, 1894; Clark, 1901, 1913, 1920;
Perrier, 1902; Grieg, 1921; Hansen, 1967, 1975) and specialists (Pawson, Sibuet, Gebruk, Massin, pers. comm.)
is Oneirophanta mutabilis (Théel, 1879), family Deimatidae and Order Elasipodida. This cosmopolitan holothurian is characteristic of the abyssal region, having been
recorded at depths between 1 800 m and 6 000 m in
all explored parts of ocean depths. This taxon could be
Oneirophanta mutabilis affinis (Ludwig, 1894), since that
is peculiar to the region studied, in the tropical eastern
Pacific Ocean, where it replaces the cosmopolitan Oneirophanta mutabilis mutabilis (Théel, 1879). However,
HOL 4 is larger than O. mutabilis affinis, and from the
description, it appears closer to O. mutabilis mutabilis.
Less similar, Oneirophanta setigera (Ludwig, 1893) has
been collected in the south-western Pacific Ocean (appendix 3).
N
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Fig. 25. Photograph taken by the ‘Epaulard’ at the
NIXO 45 site: mixed nodule-facies (C and B), holothurian
Orphnurgus sp. (HOL 3)
© Ifremer
Fig. 26. Photograph taken by the ‘Epaulard’ at the NIXO
45 site: nodule-facies C, holothurian Orphnurgus sp.
(HOL 3) © Ifremer
Fig. 27. Photograph taken by the ‘Epaulard’ at the NIXO 45 site:
nodule-facies C, holothurian Oneirophanta mutabilis (HOL 4)
© Ifremer
Fig. 28. Photograph taken by the ‘Nautile’ at the
NIXO 45 site: rock outcrop, holothurian Oneirophanta
mutabilis (HOL 4)
© Ifremer
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HOL 5
HOL 6
This white holothurian is smaller, thickset and ovoid
(figure 29), measuring a maximum of 15 cm long by 5
cm wide. Its papillae are less numerous, thicker and rigid. There are 5 to 16 pairs of dorsal papillae arranged
in a double row and 3 to 7 pairs of lateral papillae. From
observations in the study area, they are always found
on the ocean bed, in areas with or without nodules, and
consequently they may adopt a non-selective detritivorous feeding behaviour. The trail left by their passage
does not indicate that medio-ventral podia are absent.
This holothurian measures 12 to 25 cm in length and 3 to
5 cm in width (figures 30 & 31). It appears semicircular
and flattened, with the two extremities rounded. One can
see on the dorsal surface a median groove and two rows
of regularly distributed dorso-lateral papillae. The tegument appears soft and thick, and bluish or pale mauve in
colour in photographs taken by the “Nautile” (figure 27).
This taxon is found on all types of substratum, with or
without nodules, as well as on rocky surfaces, but never
in the water column.
Pawson & Foell (1983) identified this holothurian from
the DOMES C site, and it also appears in photographs
from the Peruvian Basin (Bluhm, 1994; Bluhm & Gebruk,
1999) and the Bay of Biscay (Sibuet et al., 1980).
The proposed identification, with reference to the literature (Théel, 1882, 1886; Ludwig, 1894; Perrier, 1896;
Clark, 1920; Hansen, 1956, 1975) and to specialists, is a
holothurian of the genus Benthodytes, family Psychropotidae and Order Elasipodida, referred to here as Benthodytes sp. type 1. Visible characteristics of the family
are the medio-ventral podia and a body laterally fringed
by anastomosing podia. Some members of the genus
Benthodytes have been observed swimming (Pérès,
1965; Heezen & Hollister, 1971). Pawson & Foell (1983)
identified several types of Benthodytes at the DOMES
C site. In photographs from the Peruvian Basin, Bluhm
& Gebruk (1999) identified a holothurian that was similar but very dark in colour and with some perpendicular
dorsal papillae as Benthodytes sp. type 2.
The identification proposed in accordance with the literature (Théel, 1879, 1882; Ludwig, 1894; Sluiter, 1901;
Koehler et Vaney, 1905; Hérouard, 1923; Deichmann
1930; Hansen, 1967, 1975) and in agreement with specialists is Deima validum (Théel, 1879), family Deimatidae,
Order Elasipodida. However, one would infer from the
tracks created by HOL 5 that Deima validum is not characterised by an absence of medio-ventral podia, which
contradicts the description of the podial sole made by
Hansen (1975). According to him, Deima validum does
not have medio-ventral podia, or at the very most, has
one pair reduced to pre-anal podia. More precisely, HOL
5 may be the cosmopolitan D. validum pacificum (Ludwig, 1894) collected in the tropical north-easern Pacific
Ocean between 1 618 m and 2 487 m on various substrata (appendix 3). According to Sibuet et al. (1984), Deima
validum is not a selective detritus feeder.
Data on Benthodytes species collected near to or in the
Clarion-Clipperton fracture zone are shown in appendix 3.
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Fig. 29. Photograph taken by the ‘Epaulard’ at the
NIXO 45 site: nodule-facies C, holothurian Deima
validum (HOL 5) © Ifremer
Fig. 30. Photograph taken by the ‘Nautile’ during the
NIXONAUT cruise: holothurian (collected) Benthodytes sp.
type 1 (HOL 6) © Ifremer
Fig. 31. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
rocky facies, holothurian Benthodytes
sp. type 1 (HOL 6)
© Ifremer
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HOL 7
This taxon has a cylindrical body with rounded extremities (figures 32, 33). It measures about 20 cm long and
6 cm wide. The whitish tegument is often covered by
sediment or detritus, giving the appearance of a smooth
surface. The mouth is ventral but the podia are not visible in the photographs. The distinctive feature of this
taxon is the vertical notch situated at the posterior end
of the body. This holothurian is invariably found on all
facies with nodules and sediments but never in the water column.
The proposed identification, in agreement with specialists and the literature (Ludwig, 1894; Madsen, 1953) is
Pseudostichopus mollis (Théel, 1886), family Synallactidae, Order Aspidochirotida.
This cosmopolitan holothurian has already been recorded from the Clarion-Clipperton fracture zone (Pawson
& Foell, 1983), as well as elsewhere in the north-eastern
and south-eastern Pacific, the south-eastern Atlantic and
the Indian Oceans, at depths varying between 100 m and
5 203 m on a great diversity of substrata (appendix 3).
This taxon does not appear in photographs from the Peruvian Basin (Bluhm, 1994; Bluhm & Gebruk, 1999).
As indicated by its spade-like tentacles, Pseudostichopus mollis adopts a distinctive feeding behaviour, raking
up a thin film of superficial sediments (Massin, 1982).
HOL 8
This holothurian is 10 to 20 cm long and up to 5 cm wide
(figure 24). It has a sub cylindrical or flattened and elongated form, wider than high, and with a marginal fringe. The
dorsal surface of the tegument bears numerous small irregular papillae that trap epibenthic detritus (e.g. debris
from nodules, Foraminifera, shells of pteropods etc.),
and in consequence, its colour is similar to that of this
soft substratum. Its trail is more accentuated on purely
sedimentary facies where it appears half buried. It does
not appear to be mesopelagic, always being found at the
surface of sediments.
macdonaldi (Ludwig, 1894; Sluiter, 1901), as were those
reported from the central eastern Pacific Ocean between
1 644 m and 2 149 m (appendix 3).
HOL 9
This holothurian measures up to 25 cm long and 5-6 cm
wide and has a pointed structure at one end of the body
(figure 25). Its body is bulging and cylindrical and has podia scattered all over. Those situated on the ventro-lateral
surfaces are larger and could be seen in some photographs. This holothurian has a whitish tinge, and is brown
to mauve in photographs in the rare instances when not
covered by sediment. It is only seen on nodular substrata
where it leaves a track that is clearly less deep than that
of HOL 8. It is known to be a non-selective detritus feeder,
and does not appear to be mesopelagic, never having
been seen in the water column.
This holothurian has also been reported from the DOMES
C site (Pawson & Foell, 1983), in the Peruvian Basin (Bluhm, 1994; Bluhm & Gebruk, 1999) and in the Atlantic, in
the Bay of Biscay (Sibuet et al., 1980).
Literature references (Fisher, 1907) and the advice of specialists put this holothurian in the cosmopolitan species
Mesothuria murrayi (Théel, 1886), family Synallactidae,
Order Aspidochirotida. This taxon has been recorded
from the western and eastern Pacific Ocean, as well as in
the Indian Ocean at depths varying between 725 m and
4 064 m. It was not cited in the inventories of Kaufmann &
Smith (1997) and Lauerman et al. (1996).
According to Heezen & Hollister (1971), Lemche et al.
(1976) and Billet (1991) some Mesothuria and Pseudostichopus hollow out tracks about 1 cm deep by alternately extending and contracting the whole body, since the
ventral podia are scarcely developed. These tracks are
the most abundant form of bioturbation that disturbs the
sedimentary surface to a significant degree in the depths
of the Venezuelan Basin (Young et al., 1985), the Bay of
Biscay (Mauviel & Sibuet, 1985) and the abyssal plain of
Madeira (Hugget, 1987).
This holothurian has also been reported at the DOMES C
site (Pawson & Foell, 1983), in the Peruvian Basin (Bluhm,
1994; Bluhm et Gebruk, 1999) and in the Atlantic Ocean, in
the Bay of Biscay (Sibuet et al., 1980).
The proposed identification, after reference to the literature (Lemche et al., 1976) and in the opinion of the specialists, is a species of Meseres (Ludwig, 1894), family Synallactidae and Order Aspidochirotida. It could be Meseres
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Fig. 32. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: mixed
nodule-facies (C and B), holothurian
Pseudostichopus mollis (HOL 7)
© Ifremer
Fig. 33. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodulefacies C, holothurian Pseudostichopus
mollis (HOL 7)
© Ifremer
Fig. 34. Photograph taken by the
‘Epaulard’ at the NIXO 45 site:
nodule-facies O, holothurian Meseres
macdonaldi (HOL 8) © Ifremer
Fig. 35. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise: mixed
nodule-facies (C and B), tumulus with
holothurian Mesothuria murrayi (HOL 9)
© Ifremer
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HOL 10
HOL 11
This taxon measures 20 to 25 cm long and 5 to 7 cm
wide (figures 36, 37). It appears translucid and bears a
long tapering projection on top of its anterior end. Its
ventro-lateral podia are easily distinguished, few in
number, relatively well spaced and independent, giving
the impression of raising the body. The loop formed by
the intestinal tract can be distinguished through the dorsal translucid septum, and at the anterior end of the animal is a region that reflects light. This is situated exactly
at the level of the peripharyngeal cavity where the radiating conjuncto-muscular flanges are encrusted with
calcareous bodies capable of reflecting light.
This holothurian, 17 to 20 cm long and 5 to 8 cm wide,
resembles the previous taxon in its translucent texture
through which one can see the loop of the intestinal
tract, and a white mark, which reflects the light, situated
at the peripharyngeal level (figure 38). Nevertheless, it
appears flatter and bears marginal fringes that hide the
ventro-lateral podia. A short velum in the nuchal part is
formed by the fusion of the dorsal anterior papillae, and
seems to unite four or five short projections. HOL 11 has
been observed on all types of substrata but never in the
water column.
The holothurian HOL 10 has been found on all types of
substratum, including sedimentary rocky, and with polymetallic nodules. It does not appear to be mesopelagic,
never having been seen in the water column. It has not
been observed in the Peruvian Basin nor the Bay of Biscay though forms morphologically similar to Peniagone
sp. have been identified (Sibuet et al., 1980; Bluhm, 1994;
Kaufman & Smith, 1997; Bluhm & Gebruk, 1999).
The proposed identification after referring to the literature (Hansen, 1975) and based on the opinion of specialists
is Peniagone papillata (Hansen, 1975), family Elpidiidae,
Order Elasipodida. This taxon has been collected particularly in the Clarion-Clipperton fracture zone at 3570
m. Its external characteristics are an elongated body, a
soft, whitish tegument, eight to nine pairs of podia decreasing in size towards the posterior and situated in the
posterior 2/3 of the ventral sole. It possesses four or five
pairs of dorsal papillae of which the first and sometimes
the second pair are independent, long and slender.
This taxon resembles a pink-tinted translucent species
from the abyssal depths of the Peruvian Basin, identified
as Peniagone gracilis (Bluhm & Gebruk, 1999). However,
it was characterised by having podia only in the posterior half or 2/3rds of the body.
The independent movement of the ventro-lateral podia
of HOL 10 is remarkable and recalls the model of locomotion proposed by Hansen (1972) for Scotoplanes globosa (Théel, 1879), another member of the Elpidiidae.
Its podia communicate with wide water-filled cavities
in the tegument, and local constriction of the latter, by
peristaltic movements of the epidermal muscles, allows
the independent movement of the podia. This suggests
an adaptation to locomotion on soft substrata in abyssal
environments (Hansen, 1975).
Lauerman et al. (1996) and Kaufman & Smith (1997) recorded this taxon at their station further north at 4100 m,
but it has not been found in the Peruvian Basin (Bluhm
& Gebruk, 1999), nor does it appear to be present in the
Bay of Biscay (Sibuet et al., 1980).
We propose as a hypothesis of identification a member
of either the genus Amperima (Pawson, 1965) or Peniagone (Théel, 1882), family Elpidiidae and Order Elasipodida. HOL 11 and the following taxa (HOL 12 to HOL 15)
share the characteristics of the groups Peniagone and
Amperima (and part of the group Ellipinion), that is, an
ovoid body, a wide velum consisting of two pairs of papillae, of which the median ones are generally longer
than the laterals, podia bordering the posterior half or
third of the body or distributed uniformly along the two
lateral parts of the body.
Identification can not be taken further without a detailed
study of the spicules. However, based on the literature
(Théel, 1882; Ludwig, 1894; Clark, 1920; Hansen, 1975),
one could compare this taxon with Peniagone vitrea
(Théel, 1882) which has been collected in the region
(between 10°N-10°S, cf. appendix 3). The external characteristics of this holothurian are an elongated body, a
wide velum that can be folded either towards the anterior
or the posterior, and 6 to 9 pairs of podia that decrease
in size towards the posterior and border the posterior
third of the ventral sole. As in most of the Elasipods,
members of the genera Amperima and Peniagone comb
a superficial film of sediments leaving few traces of their
passage (Massin, 1982).
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Fig. 36. Photograph taken by the
‘Nautile’ at the NIXO 45 site: cliff, holothurian
Peniagone papillata (HOL 10)
© Ifremer
Fig. 37. Photograph taken by
the ‘Epaulard’ at the NIXO 45
site: mixed nodule-facies B and
plates: holothurian Peniagone
papillata (HOL 10)
© Ifremer
Fig. 38. Photograph taken by
‘E.D.1’ during the Copano cruise:
nodule-facies B, holothurian
Peniagone vitrea (HOL 11)
© Ifremer
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HOL 12
This taxon only differs from the preceding one by its velum, which is formed by two small anterio-dorsal projections (figure 39). It can reach 20 cm in length and 8 cm
width, and is found on all substrata but never in the water
column.
Pawson & Foell (1983) found this holothurian at the DOMES C site in the Clarion-Clipperton fracture zone. Lauerman et al. (1996) and Kaufman & Smith (1997) have also
identified it at 4100 m at a station further north and close
to the central part of the Californian coast. This taxon has
also been found in the abyssal depths of the Peruvian Basin (Bluhm & Gebruk, 1999) where it appears translucid
brown. In contrast, no similar holothurian has been found
in the Bay of Biscay (Sibuet et al., 1980).
Based on the literature (Ludwig, 1894; Hansen, 1975), the
proposed identification is a member of the genus Amperima (Pawson, 1965) or Peniagone (Théel, 1882), family Elpidiidae and Order Elasipodida, or Achlyonice sp. (Théel,
1879). Achlyonice ecalcarea has been collected between
2 780 m and 4 924 m in the Pacific Ocean (appendix 3),
and therefore we suggest this latter identification.
HOL 13
This holothurian is distinguished from the preceding
ones by the velum, which is formed of two pairs of dorsal papillae divided into two long projections, two other
shorter ones arranged laterally, and a buccal orifice situated on a retractable tube (figures 40, 41). Its body is
broader, ovoid and measures up to 30 cm long and 10
cm wide. Its ventral face is bordered by nine or ten pairs
of podia decreasing in size towards the posterior.
This taxon has been observed on all types of substrata
but never in the water column. It has been identified in
photographs from the Peruvian Basin and Bay of Biscay.
Pawson & Foell (1983) identified it from the DOMES C
site, and Kaufman & Smith (1997) have also identified
it from their site further north. However, it has not been
seen in the Bay of Biscay (Sibuet et al., 1980).
As in the case of HOL 10, this taxon is similar to a translucent, pink holothurian from the abyssal depths of the
Peruvian Basin (Bluhm & Gebruk, 1999). However, this
latter taxon, which has been identified as Peniagone gracilis, is narrower and its podia are not restricted to the
posterior half or 2/3 of the body.
The proposed identification, based on the literature
(Théel, 1882; Perrier, 1896, 1902; Hérouard, 1923; Agatep, 1967; Hansen, 1975) and the advice of specialists,
is Amperima rosea (Perrier, 1896), family Elpidiidae and
Order Elasipodida. However, this taxon has never been
collected in the Pacific Ocean and has only been found
between the Azores and Portugal at depths between
4 060 m and 5 000 m (cf. appendix 3).
HOL 14
This holothurian reaches a maximum of 25 cm long and
5 cm wide (figure 42). It only differs from the preceding
ones by being more massive and by its velum, which is
formed of four wide, equal-sized, anterio-dorsal projections that can be folded either backwards over the body
or forwards towards the substratum. It has been observed on all types of substratum but never in the water
column, and thus it does not appear to be mesopelagic.
This taxon has perhaps been found in photographs from
the region of the DOMES sites, recorded as a Peniagone
sp., either type 1 or type 2 (Pawson & Foell, 1983). Lauerman et al. (1996) and Kaufman & Smith (1997) did not
cite it in their faunistic inventories, nor has it been reported from the Bay of Biscay (Sibuet et al., 1980).
The proposed identification, based on the literature (Ludwig, 1894; Clark, 1920b; Hansen, 1975; Madsen, 1953)
and in agreement with the specialists, is Peniagone purpurea (Théel, 1882), as in the case of specimens collected in the Indo-Pacific region (cf. appendix 3). Peniagone
purpurea has the following external characteristics: a
thickset body with seven to nine pairs of podia, the first
two being independent and ventral while the other five
to seven pairs are close together and partly fused. The
velum consists of two pairs of very long, thin papillae
joined together at the base. The edge of the velum is prolonged in a lateral fringe covering the podia.
In the figures, the velum sometimes appears folded
towards the anterior with the extremities emerging on
the substratum, or sometimes it is held high, almost
perpendicularly. This could have various functions, sensorial and perhaps even respiratory, by increasing the
surface area of the body and facilitating the exchange
of fluids in the water-vascular system by its movements
(Hansen, 1975).
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Fig. 39. Photograph taken by the ‘Epaulard’
at the NIXO 45 site: mixed nodule-facies C,
holothurian Achlyonice ecalcarea (HOL 12)
© Ifremer
Fig. 40. Photograph taken by the ‘E.D.1’
device during the Copano cruise: nodule-facies
C, holothurian Amperima rosea (HOL 13)
© Ifremer
Fig. 41. Photograph taken by a ‘Troïka’ during
the Copano 1 cruise: nodule-facies C, holothurian
Amperima rosea (HOL 13)
© Ifremer
Fig. 42. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
nodule-facies O, holothurian Peniagone purpurea
(HOL 14) © Ifremer
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HOL 15
This whitish taxon, maximum dimensions 20 cm long
and 5 cm wide, seems to be quite rare in the study zone
(figures 43, 44). It does not appear to have a preference
in relation to substratum, occurring on nodules or sediments, and it does not appear to be mesopelagic.
This taxon has also been found in the Peruvian Basin
(Bluhm & Gebruk, 1999) but was not recorded among the
echinoderms in the Bay of Biscay (Sibuet et al., 1980).
Except for its two large characteristic anterio-dorsal
projections, it differs little from the other holothurians
of the genera Amperima (Pawson, 1965) or Peniagone
(Théel, 1882), family Elpidiidae and Order Elasipodida.
As a more precise hypothesis of identification, based on
the literature (Théel, 1882; Clark, 1920b; Madsen, 1953;
Hansen, 1956, 1975) and the specialists, one could propose Amperima naresi (Théel, 1882), which has been
collected in the study zone and is very similar. Its external characters are a body more ovoid than in the other
members of the group, and eight to ten pairs of broad
ventro-lateral podia, which border the whole sole, decrease in size towards the posterior, and are distributed more or less uniformly along the sides of the body.
These podia are slightly fused in the posterior part. The
velum, composed of two pairs of papillae, is well developed and can take different forms (Hansen, 1956).
Appendix 3 presents possible identifications for taxa
HOL 10 to HOL 15 by describing the holothurians recorded belonging to the genera Amperima (Pawson, 1965)
or Peniagone (Théel, 1882). Specimens of Amperima naresi would have come from between 10°S and the central eastern Pacific Ocean, between 2 010 m and 7 130 m
depth.
HOL 16
This quite remarkable holothurian measures at the very
most 40 cm long and 12 cm wide, with a very large caudal extension held perpendicular to the substratum, and
which is about the same length as the body and as wide
as the latter in its most expanded part (figures 45, 46).
The body is very flattened on the ventral surface and arched dorsally. The extremities are strongly rounded and
the whole body is bordered by a wide dark podial fringe
on the trivium. The caudal appendage is fleshy and more
indented along a median grove. This appendage originates in an upper dorsal region, about 1/4 to 1/3 of the
way from the posterior end. This holothurian can be very
dark or almost translucent, and the colour photographs
taken by the “Nautile” show a bluish tinge for the translucent specimens.
This taxon has been observed on all types of substratum
and occasionally in the water column. Pawson & Foell
(1983) identified it from the DOMES C site though Kaufman & Smith (1997) did not see it at their station further
north-east, and neither did Bluhm (1994; Bluhm & Gebruk, 1999) in the Peruvian Basin. On the other hand, it is
present in the Bay of Biscay (Sibuet et al., 1980).
The proposed identification, based on the literature
(Théel, 1882; Hérouard, 1902, 1923; Deichmann, 1940;
Madsen, 1953) and on the advice of specialists, is Psychropotes semperiana (Théel, 1882), family Psychropotidae, Order Elasipodida. This holothurian has not
yet been collected in the study zone nor elsewhere in
the Pacific, but only in the Indian and Atlantic Oceans at
depths of 3 460 m to 5 600 m.
The translucent versions of this taxon could be either
juveniles that become darker in the adult stage (Miller &
Pawson, 1990), or they could belong to another species,
Psychropotes hyalinus, collected in the Pacific north-east
at 5 891 m (Pawson, 1985). This latter species is very close to Psychropotes semperiana but differs in its transparency and the characteristics of its spicules. In addition,
it is known to be a facultative swimmer, using its large
marginal fringe as fins. Elsewhere it has been collected
at 5 m above the substratum at a depth of 5890 m in the
tropical north-easern Pacific Ocean (Pawson, 1985). In
films taken by the “Nautile”, one can see it making swimming movements when it contacts the manipulator arm,
by twisting then gliding. This could be an avoidance behaviour in case of predation.
The tracks left by this taxon are difficult to see in the
numerous photographs from the study zone. That might
suggest Psychropotes semperiana, which has relatively
small podia on the medio-ventral part of the plantar sole,
distributed in two rows of 7 to 25. The nodules appear
darker, as if cleaned by the animal’s passage.
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Fig. 43. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodule-facies
C, holothurian, Amperima naresi (HOL 15)
© Ifremer
Fig. 45. Photograph
taken by the
‘Epaulard’ at the NIXO
45 site: mixed nodule-facies (C and B), holothurian Psychropotes
semperiana (HOL)
© Ifremer
Fig. 44. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodule-facies
O, holothurian, Amperima naresi (HOL 15)
© Ifremer
Fig. 46. Photograph taken by the ‘Epaulard’ at the Nixo 45 site: nodule-facies C,
holothurian Psychropotes semperiana
(HOL 16) © Ifremer
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HOL 17
This holothurian differs from the preceding one particularly in the location of the caudal appendage, which is in
the extension of the posterior part of the body, behind
the bivium (figures 47, 48, 49, 50). This taxon reaches a
maximum of 50 cm in length and 15 cm width with a
caudal appendage up to 30 cm long. The latter appears
more fleshy and rigid in its first third, and more supple at
its extremity, which can be flexed during swimming. The
marginal fringe is much less developed than in the preceding species and demarcated from a well-developed
peristomal membrane. In some photographs, the tail appears shorter, the peristomal membrane less spreading
and the ventro-lateral podia more free and prominent.
This taxon appears very dark in black and white photographs but pale yellow to orange in the colour ones taken by the “Nautile”. In film from the NIXONAUT cruise,
it performs a swimming motion, rhythmically arching
the axis of its body after inducing a pulsation in its peristomal membrane, in opposition to movement in its caudal appendage. This taxon was only observed on ocean
floor with a dense covering of nodules.
Based on the literature (Théel, 1882; Ludwig, 1894; Clark,
1920; Madsen, 1953; Parker, 1963; Bayer, 1970; Hansen,
1975; Luke, 1982) and the advice of specialists, we propose as a hypothesis of identification, Psychropotes
longicauda (Théel, 1882), family Psychropotidae, Order
Elasipodida. This cosmopolitan holothurian has been
reported between 2 210 m and 5 203 m (appendix 3).
It bears between 7 and 25 pairs of independent podia,
18 peribuccal tentacles, a marginal fringe composed of
two to nine pairs of podia and two to eight pairs of very
small dorsal papillae. It can take on yellow to deep violet
tones, always with the plantar sole darker.
According to Khripounoff & Sibuet (1980), Psychropotes
longicauda is a detritus feeder, which selects the particles richest in assimilable compounds, and occasionally
a scavenger. It has been recorded in all the oceans at
depths between 2 210 m and 5 203 m on a great variety
of substrata. According to Mortensen (1927), the caudal
appendage is used in swimming, though Miller & Pawson (1990) stated that only the juveniles are mesopelagic
and some of these have been collected 500 m above the
benthos (Billet et al., 1985).
Pawson & Foell (1983) have identified this holothurian
at the DOMES C site, while Kaufman & Smith (1997) did
not see it at 4 100 m. Bluhm (1994) and Bluhm & Gebruk
(1999) recorded it in the Peruvian Basin and it is also present in the Bay of Biscay (Sibuet et al., 1980).
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Fig. 47. Photograph taken
by the ‘Nautile’ during the
NIXONAUT cruise: nodule-facies C, holothurian (collected)
Psychropotes longicauda
(HOL 17)
© Ifremer
Fig. 48. Photograph taken by the ‘Nautile’ during
the NIXONAUT cruise: nodule-facies C, holothurian
Psychropotes longicauda (HOL 17)
© Ifremer
Fig. 49. Photograph taken by the ‘Nautile’
during the NIXONAUT cruise: nodule-facies O,
holothurian Psychropotes longicauda (HOL 17)
© Ifremer
Fig. 50. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodule-facies C,
holothurian Psychropotes longicauda (HOL 17)
© Ifremer
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HOL 18
This swimming holothurian is dark in colour and
brown-violet in films taken by the “Nautile” (figures
51, 52, 53, 54). Its fleshy body (maximum dimensions
25 cm long and 10 cm wide), is slightly dorso-ventrally
flattened. The anterior part bears a velum combining
12 to 14 broad dorso-ventral papillae into a membrane
encircling 3/4 of the body. The anterio-ventral branch
is encircled by 20 small tentacles whose extremities
are divided in dendritic ramifications. The ventro-lateral podia are divided into two groups, one forming
two fans each with six to nine pairs of long podia joined together posteriorily, and the other with three to
six pairs of anterior, more rudimentary, podia.
Most of the time this holothurian is seen swimming
over a great variety of facies of nodules and sediments.
The tracks it leaves on the substratum are formed by
the weight of the body resting on the ends of the three
or four pairs membranous posterior podia and by the
terminal parts of the folded tentacles. When following
the swimming movements in film taken by the “Nautile”, the interval for each propulsion was estimated at
5 seconds. Miller & Pawson (1990) recorded a cycle of
7.5 seconds, during which the animal stayed vertical
and beat the water by negative rheotropism with an
undulating movement of its velum.
Pawson & Foell (1983) identified this holothurian at
the DOMES C site although Kaufman & Smith (1997)
did not find it at 4100 m at their station further northeast in the Pacific Ocean, perhaps because it has a
more mesopelagic lifestyle. Bluhm (1994) and Bluhm
& Gebruk (1999) definitely identified it in the Peruvian
Basin, and it is also present in the Bay of Biscay (Sibuet et al., 1980).
The hypothesis of identification, based on the literature (Pawson, 1976, 1982; Ohta, 1985; Miller & Pawson, 1990) and the advice of specialists, is Enypniastes
eximia (Théel, 1882), family Pelagothuriidae and Order Elasipodida. This holothurian has been observed
swimming between 2.5 m and 5 m above the substratum, except for young specimens, which have been
collected more than 3 000 m above the bottom by
Billett et al. (1985), or even at the ocean surface (Herouard, 1923). Details of its swimming were described
by Ohta (1985) and Miller & Pawson (1990). However,
in photographs and films from the Clarion-Clipperton
fracture zone, Enypniastes eximia adopts a swimming
posture typical of currents, and uses its velum as a
sail, opposed to the direction of the current, and its
ventro-lateral podia as brakes. According to in situ
observations, young E. eximia are smaller and translucent pink in colour (Billett et al., 1985; Miller & Pawson, 1990).
Enypniastes eximia is known to be cosmopolitan and
has been recorded at depths between 516 m and 5 689
m (appendix 3). This holothurian spends 10% of its time
on the substratum in order to ingest sediment in a nonselective manner (Miller & Pawson, 1990). Thus it is not
a suspension feeder and does not feed in the water
column (Ohta, 1985). However, it could be at an intermediate stage of evolutionary adaptation to an essentially mesopelagic life, as seen in Pelagothuria natatrix
(Ludwig, 1894; Chun, 1900; Etman, 1926). The latter is
presumed to be a suspension feeder since it has only
been collected in the water column (Ohta, 1985).
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Fig. 51. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodule-facies C, holothurian (swimming)
Enypniastes eximia (HOL 18)
© Ifremer
Fig. 52. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
nodule-facies C, holothurian (swimming)
Enypniastes eximia (HOL 18)
© Ifremer
Fig. 53. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodule-facies C,
holothurian Enypniastes eximia (HOL 18)
© Ifremer
Fig. 54. Photograph taken by the ‘Epaulard’
at the NIXO 45 site: nodule-facies C, holothurian
Enypniastes eximia (HOL 18)
© Ifremer
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HOL 19
This taxon with an ovoid body can measure up to 32 cm
long by 10 cm wide (figures 55, 56, 57, 58, 59, 60), and in situ,
it is bright pink. The velum is formed by four dorsal papillae
of which the central pair is longer than the laterals. At the
posterior of the body are four pairs of ventral, membranous
podia, laterally arranged. The mouth is in a retractable tube
oriented towards the posterior. About ten tentacles, branched at their ends, surround the mouth and folds mark the
dorsal surface of the body. A membranous fringe joins the
velum to the ventro-posterior podia laterally.
In photographs and film from the “Nautile”, this holothurian
is mostly seen swimming. Whether suspended vertically
and maintaining its flotation by beating its buccal tentacles,
or with its body-axis parallel to the substratum, it propels
itself by flexing its body, which is bent at the level of the
velum, in rhythmic opposition to the pulsation of its ventroposterior podia. When on the substratum, it feeds from the
superficial film covering the nodules and sediments with
the aid of its tentacles, which are forked at their ends (figure
60). The axis of the mouth and tentacles form a right angle
in relation to the axis of the squat body, held up by the two
series of ventro-posterior podia.
Pawson & Foell (1983) identified this holothurian at the DOMES C site although Kaufman & Smith (1997) did not find it,
perhaps because it lives mostly in the water column. Bluhm
(1994) and Bluhm & Gebruk (1999) did not record it in the
Peruvian Basin, though it is present in the Bay of Biscay at a
depth of 3000 m (Sibuet et al., 1980).
The hypothesis of identification based on the advice of specialists is Peniagone leander (Pawson & Foell, 1986), family
Elpidiidae, Order Elasipodida. However, the authors described this taxon on the basis of photographs, which were
taken between 14°33’N-127°39’W and at around 4 500-5 000
m depth. They referred to the quality of photographic detail
that enabled them to see all the taxonomic parts required
for its identification, and the considerable difficulty in collecting an organism that is so fragile and for the most part,
is mesopelagic.
HOL 20
simultaneously with its posterior paired projections and its
10 peribuccal, branched tentacles.
Pawson & Foell (1983) identified this holothurian at the DOMES C site although Kaufman & Smith (1997) did not record
it at 4100 m at the station further north-east, perhaps because it lives more in the water column. Bluhm (1994; Bluhm
& Gebruk, 1999) recorded it in the Peruvian Basin, and it
is known from photographs from the deep north Atlantic
Ocean, in the Bay of Biscay (Sibuet et al., 1980).
The hypothesis of identification proposed on the basis of
literature (Théel, 1882; Herouard, 1899, 1902, 1923; Madsen,
1953; Barnes et al., 1976) and the advice of specialists is Peniagone diaphana (Théel, 1882), family Elpidiidae, Order
Elasipodida. This species has been collected in the Pacific,
Atlantic and Indian Oceans at depths between 1 520 m and
5 600 m (appendix 3). Peniagone diaphana has been observed from a submersible (Barnes et al., 1976) in such a way
that its rhythmic swimming pulsations could be determined
as 10 to 20 cycles per minute. It is known that it can maintain
itself vertically in the water column for more than 12 hours
and it was once found at more than 70 m above the bottom
surface. According to Hansen (1975), the specific weight of
this pelagic species is close to that of water, and it utilises
currents as a means of displacement.
An interesting hypothesis for the function of its swimming
movements has been proposed by Barnes et al. (1976).
They suggest that they facilitate respiration and cause a
current that puts particles of detritus from the water-sediment boundary into suspension, and which thus feed the
animal. The hypothesis was proposed because Peniagone
diaphana seems to be essentially mesopelagic, having
been observed from a submersible to spend more than 12
hours continuously in the water column. It seems to be a
suprabenthic detritus feeder on the basis of the contents of
its digestive tract, which include the tests of Foraminifera,
spicules of sponges and holothurians, as well as other detritus of benthic origin. Barnes et al. (1976) attributed a special
feeding behaviour to it, that of a detritus feeder with the lifestyle almost of an agile suspension feeder. However, just as
in Peniagone leander, this taxon may perhaps sometimes
settle on the substratum in order to feed itself, like all detritus feeders, and the long time spent in the water column
could perhaps be an artefact caused by the presence of the
submersible disturbing its usual way of life.
This holothurian closely resembles the preceding one in its
body shape and its mesopelagic life (figure 61). However, it
presents the following characters: a flattened body, 15 cm
long and 7 cm wide, a translucid tegument, through which
the pigmented loop of the digestive tract is visible, a broad
velum, formed of four dorsal papillae and directed forwards
and prolonging the dorsal surface, four to five pairs of fused
podia arranged in two posterior winglike projections Since
it has never been seen on substratum, only its position in
the water column can be described, and here it swims by
maintaining its body axis vertically and beating the water
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Fig. 55. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
nodule-facies C, holothurian (swimming)
Peniagone leander (HOL 19)
© Ifremer
Fig. 56. Photograph taken by the ‘Epau-
lard’ at the NIXO 45 site: nodule-facies C,
holothurian (swimming) Peniagone leander
(HOL 19)
© Ifremer
Fig. 58. Photograph taken by a ‘Troïka’ during the
Copano 1 cruise: nodule-facies C, holothurian (swimming)
Peniagone leander (HOL 19) © Ifremer
Fig. 57. Photograph taken
by the ‘Epaulard’ at the
NIXO 45 site: nodule-facies
C, holothurian (swimming)
Peniagone leander (HOL 19)
© Ifremer
Fig. 59. Photograph taken by the
‘Nautile’ during the NIXONAUT
cruise: nodule-facies C, holothurian
on the ocean floor Peniagone
leander (HOL 19)
© Ifremer
Fig. 60. Photograph taken by
the ‘Nautile’ during the NIXONAUT cruise:
nodule-facies C, holothurian (swimming)
Peniagone diaphana (HOL 20)
© Ifremer
ENGLISH ATLAS PHOTO pages.indd 35
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HOL 21
This holothurian measures up to 55 cm long and 5 cm
wide and its body is cylindrical to fusiform (figures 61,
62, 63, 64, 65, 66). In colour it is dark purple or violet
with paler patches. The dorsal papillae form an irregular
fringe around the anterior dorsal extremity. The oral region bears tentacles and is directed ventrally. Two rows
of small more or less protruding papillae are aligned on
the dorsal surface. The posterior ends in a blunt point,
sometimes with some short tapering projections. The
non-gelatinous tegument resembles leather. This holothurian appears entirely benthic since it has never been
reported from the water column. It has been found on
different types of nodule substrata and purely sedimentary ones. Its tracks on sediments are characterised by a
double row of relatively large and well-spaced ventro-lateral podia, and by an absence of medio-ventral podia.
ventories of Kaufman & Smith (1997) or Lauerman et al.
(1996). Bluhm (1994) and Bluhm & Gebruk (1999) found
it in the Peruvian Basin and it has been recognised in
photographs from the Bay of Biscay in the north Atlantic
Ocean (Sibuet et al., 1980).
The hypothesis of identification in agreement with Pawson (pers. comm.) is Psychronaetes hanseni (Pawson,
1983), family Laetmogonidae, Order Elasipodida, which
has been collected in the central Pacific Ocean and especially in the study zone, between 4 800 m and 5 200 m
(appendix 3). Externally, the holotype does not have peribuccal papillae but has 15 tentacles with broad oral discs
and small short stalks. The 15 ventro-lateral podia per row
are approximately 10 mm long and 8 mm wide, triangular
in shape and evenly distributed. The dorsal papillae are
arranged in two rows with 30 in each row. At the anterior
end, about 15 papillae, measuring 15 mm long by 9 mm
wide for the largest, form an irregular fringe.
Pawson & Foell (1983) identified this holothurian from
the DOMES C site although it was not recorded in the in-
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Fig. 61. Photograph taken by the
‘Nautile’ during the NIXONAUT cruise:
capture of a holothurian Psychronaetes
hanseni (HOL 21)
© Ifremer
Fig. 62. Photograph taken by ‘Deep
Tow’ at the Echo I site: nodule-facies
C, holothurian Psychronaetes hanseni
(HOL 21)
© Ifremer
Fig. 63. Photograph taken by the ‘Epaulard’ at
the NIXO 45 site: nodule-facies C, holothurians
Psychronaetes hanseni (HOL 21) and
Oneirophanta mutabilis (HOL 4)
© Ifremer
Fig. 64. Photograph taken by the ‘Nautile’ during the NIXONAUT cruise:
nodule-facies O, holothurian Psychronaetes hanseni (HOL 21) © Ifremer
Fig. 66. Photograph taken by the ‘Nautile’
during the NIXONAUT cruise: on ancient dredge
tracks, two holothurians Psychronaetes hanseni
(HOL 21) and Mesothuria murrayi (HOL 9)
© Ifremer
Fig. 65. Photograph
taken by the ‘Nautile’ during the NIXONAUT cruise:
nodule-facies C, holothurian Psychronaetes hanseni
(HOL 21) © Ifremer
ENGLISH ATLAS PHOTO pages.indd 37
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HOL 22
This holothurian can reach up to 35 cm in length and 12
cm width (figure 67). It has a very dark tegument and in
consequence, the detail of the dorsal surface is very difficult to discern. In some photographs, however, a double row or bands of dorsal papillae can be seen, as well
as a marginal fringe bordering the sides of the body. In
addition, it is characterised by the absence of a distinct
anterior region, as was the case for the preceding taxon.
HOL 22 is the only holothurian observed in the study
zone that leaves a trail free of nodules on substrata with
a dense covering of nodules. This gives an idea of its
mass and power and the perturbation caused in the edaphic environment by its passage. It has never been seen
in the water column.
Pawson & Foell (1983) identified similar holothurians
from the DOMES C site, though Kaufman & Smith (1997)
and Lauerman et al. (1996) did not record it. Bluhm (1994)
and Bluhm & Gebruk (1999) reported comparable taxa in
the Peruvian Basin, and it is also present in photographs
from the Bay of Biscay in the northern Atlantic Ocean
(Sibuet et al., 1980).
The hypothesis of identification proposed by reference
to the literature (Théel, 1879, 1882, 1886a, 1886b; Ludwig, 1894; Madsen, 1953; Hansen, 1975; Luke, 1982) and
specialists is a member of the Order Elasipodida and
family Laetmogonidae, Laetmogone sp. Holothurians
identified as Laetmogone wyville thomsoni (Theel, 1879;
Madsen, 1953) have been collected in the north-east and
south-east Pacific Ocean (cf. appendix 3).
HOL 23
This holothurian measures up to 30 cm long and 6 cm
wide (figures 68, 69, 70, 71). Its dark body is semi-cylindrical, flattened on the ventral surface and supported by
a broad podial fringe. In black and white photographs
it appears dark, and in colour photographs, pale violetbrown on the dorsal surface with much darker shades
on the ventral plantar surface. In figure 69, one can distinguish two dorsal tapering papillae perpendicular to
the axis of the body. One small dorsal apophysis is situated approximately 1/4 of the way from the posterior end.
The broad podial fringe is used in swimming, putting the
holothurian in a medio-transverse plane while keeping
the dorsal apophysis upright (figure 71). This holothurian has been observed on all types of ocean floor and
its tracks are easily discernible (figure 70).
Pawson & Foell (1983) identified it from the DOMES
C site though it has not been recorded by Kaufman &
Smith (1997) or Lauerman et al. (1996) at the station further north-east at 4 100 m. Bluhm (1994) and Bluhm &
Gebruk (1999) also found it in the Peruvian Basin, but
it has not been seen in the Bay of Biscay in the north
Atlantic Ocean (Sibuet et al., 1980).
The proposed identification, based on the literature
(Ludwig, 1894; Koehler et Vaney, 1905; Clark, 1920; Madsen, 1953; Hansen, 1956, 1975) and the opinion of specialists, is Psychropotes verrucosa (Ludwig, 1894), family
Psychropotidae and Order Elasipodida. This holothurian
exhibits the external characters that can be seen in some
photographs, such as 15 or 16 peribuccal tentacles, one
to four pairs of small dorsal papillae, a conical and retractable dorsal appendage, a marginal fringe composed of ventro-lateral podia and their water-vascular
system, and a tegument covered with excrescences that
each contain a giant spicule in the form of a cross; in
older individuals, the tegument has longitudinal bands
of transverse ridges. One can explain the absence of definite tracks on the substratum by the fact that this taxon
only has medio-ventral podia (56 pairs).
This species has been reported from many localities
from the Indian to the eastern Pacific Ocean at depths
varying between 2 417 m and 7 250 m (appendix 3), but
never in the water column.
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Fig. 67. Photograph taken by the ‘Epaulard’ at the
NIXO 45 site: nodule-facies C: holothurian
Laetmogone sp. (HOL 22)
© Ifremer
Fig.68. Photograph taken by the ‘Epaulard’ at the NIXO 45 site:
nodule-facies O, holothurian Psychropotes verrucosa (HOL 23)
© Ifremer
Fig. 70. Photograph taken by the ‘Epaulard’ at
the NIXO 45 site: nodule-facies C sparse,
holothurian Psychropotes verrucosa (HOL 23)
© Ifremer
Fig. 69. Photograph
taken by the ‘Epaulard’ at
the NIXO 45 site: mixed
nodule-facies (C and B),
holothurian Psychropotes
verrucosa (HOL 23)
© Ifremer
Fig. 71. Photograph taken by the ‘Nautile’ at the NIXO 45 site: nodule-facies O,
holothurian (swimming) Psychropotes verrucosa (HOL 23) © Ifremer
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HOL 24
This holothurian measures 20 to 30 cm long and 5 to
7 cm wide (figure 72). Its very dark, smooth tegument
reflects light, as though armoured and marked by transverse folds. A relatively thin podial fringe encircles the
body. The anterior region is very distinct and almost
hexahedral in shape. It occurs on sedimentary facies and
rock outcrops but has never been observed in the water
column, and its tracks can not be discerned. Outside the
study zone, it has not been seen in photographs from
the Peruvian Basin nor those from the Bay of Biscay.
The proposed identification is a member of the family
Psychropotidae, Order Elasipodida. The identification
can not be taken further, either by reference to the literature (Sluiter, 1901; Heding, 1940; Hansen, 1975) or to specialists, because this species was previously unknown
to science. It has certain characteristics in common with
Benthodytes sibogae (Sluiter, 1901; Hansen, 1975) that
has been collected around Indonesia between 694 m
and 2 798 m.
HOL 25
This taxon is about 30 cm long and 10 cm wide, very
flattened and dark, and is found on all types of sedimentary facies and with nodules (figures 73, 74). The podial
margin is developed and continuous with the widely
spreading peribuccal tentacles. Unfortunately the dark
dull colour of the tegument prevents any morphological
detail from being distinguished, and the tracks left by its
passage could not be seen either. One individual of HOL
25 was filmed on a voyage during the NIXONAUT cruise,
raising itself half way up the side of a cliff and then diving, flattened, with the aid of its broadly spread fringe
in order to alight several metres lower. This was perhaps
a reaction to the approach of the “Nautile” or done under the influence of its hydrodynamic turbulence.
The hypothesis of identification based on the literature
(Théel, 1879, 1882, 1886a, 1886b; Ludwig, 1894; Madsen,
1953; Hansen, 1975; Luke, 1982) and specialists is a member of the Order Aspidochirotida, family Synallactidae,
Paelopatides sp. (Théel, 1886). Paleopatides confundens
(Théel, 1886) has been recorded in the north and southeast Pacific Ocean, especially in the study zone between
450 m and 4 070 m and P. suspecta at 2 323 m at 8°N in
the Pacific (appendix 3).
Pawson & Foell (1983) identified Paleopatides species
of similar appearance at the DOMES C site. Kaufman
& Smith (1997) and Lauerman et al. (1996) did not find
this genus at their station. Bluhm (1994) and Bluhm &
Gebruk (1999) have also recorded Paleopatides in the
Peruvian Basin that were brown-orange in colour and
had a somewhat convex dorsal surface and two rows
of short papillae. Similar taxa have been reported from
the Bay of Biscay in the northern Atlantic Ocean (Sibuet
et al., 1980).
According to Gage et al. (1985), holothurians identified
as Paelopatides sp. have been photographed in the
north-east Atlantic Ocean at 1942-1949 m and at 1100 m
in the north-east Pacific Ocean. They appeared to flee
with a sinusoidal motion at the approach of the trawl.
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Fig. 72. Photograph taken by the
‘R.A.I.E.’ at the NIXO 42 site: mixed
nodule-facies O and rock, holothurian
Benthodytes sibogae (HOL 24)
© Ifremer
Fig. 73. Photograph taken by the
‘Epaulard’ at the NIXO 45 site: nodulefacies O, holothurian Paelopatides
confundens (HOL 25)
© Ifremer
Fig. 74. Photograph taken by
the ‘Epaulard’ at the NIXO 45
site: nodule-facies O, holothurian Paelopatides sp. (HOL 25)
© Ifremer
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HOL 26
This holothurian measures 16 to 20 cm in length and 7
cm width (figure 75). It is very flattened, paler in colour
than HOL 25 and with the edges and the dorsal median
groove darker. The extremities of the body are rounded.
One distinctive external character is the presence of
some long tapering dorsal papillae. This holothurian has
only been observed on surfaces with nodules, where
its tracks are indistinct, and never in the water column,
although it has a morphology suitable for vertical movement due to the presence of a broad podial fringe
(Hansen, 1975).
Pawson & Foell (1983) identified similar taxa from the
DOMES C site. Kaufman & Smith (1997) and Lauerman
et al. (1996) did not find this genus at their station, and
neither did Bluhm (1994; Bluhm & Gebruk, 1999) in inventories from the Peruvian Basin. This taxon also seems
to be absent from the Bay of Biscay in the north Atlantic
Ocean (Sibuet et al., 1980).
The identification proposed, based on the literature
(Théel, 1882; Ludwig, 1894; Sluiter, 1901; Edwards, 1907;
Fisher, 1907; Mitsukuri, 1912; Clark, 1913, 1920; Oshima,
1915, 1916, 1919; Parker, 1963; Hansen, 1975) and the advice of specialists, is Pannychia moseleyi (Théel, 1882),
family Laetmogonidae. It has been recorded in the north,
south-east and south-west Pacific Ocean at depths varying between 212 m and 2 599 m (appendix 3). External
characters of this taxon are 20 tentacles, ventro-lateral
podia in a double row to the extremities (34 to 45 podia
per row), which terminate in discs in the form of suckers, 40 smaller medio-ventral podia, numerous small
papillae distributed over all the body, and 10-20 longer
papillae arranged bilaterally on the dorsal radii.
pale or sometimes off-white dorsal surface that has a
speckled appearance. It has never been seen swimming
although its morphology would be suitable. Östergren
(1938) suggested that this bicoloured pattern could be
an aid to escaping predation by swimming, as in numerous Elasipods. This taxon has only been observed on
facies with nodules and it does not leave any visible trail
in photographs. A similar taxon appears in the photographs from the Bay of Biscay but it does not show the
bicoloured pattern seen in the holotype HOL 27.
The identification proposed by reference to the literature
(Théel, 1882, 1886; von Marenzeller, 1893; Ludwig, 1894;
Perrier, 1896, 1902; Koehler et Vaney, 1905; Clark, 1913,
1920, 1923a, 1923b; Oshima, 1915, 1916, 1919; Grieg, 1921;
Hérouard, 1923; Deichmann, 1930, 1940, 1954; Heding,
1940; Madsen, 1953; Hansen, 1975) and on the advice of
specialists is a holothurian of the genus Benthodytes, family Psychropotidae, Order Elasipodida. It most closely
resembles Benthodytes typica (Théel, 1882) which has
been reported between 315 m and 4 700 m between 8°N
and 6°S in the Pacific Ocean (appendix 3). The external
characters of Benthodytes typica are, among others, 15
to 20 peribuccal tentacles, circum-oral papillae and well
pigmented water-vascular system on the ventro-lateral
podia that is equally visible from the dorsal and ventral surfaces. The dorsal papillae are few (three or four
pairs), small and aligned in two rows along the anterior
part of the dorsal radii. The tegument is soft and mucosal for most of the time. The speckled appearance of the
dorsal tegument comes from the fact that it is covered
with small mauve swellings on a dark back ground. The
ventral surface is uniformly dark violet.
HOL 27
This taxon is fairly cosmopolitan having been reported at
depths varying between 315 m and 4 700 m. According
to Miller & Pawson (1990), specimens have been collected in the Atlantic Ocean up to an altitude of 3 400 m
above the bottom, indicating that only the juvenile forms
are bentho-pelagic (Grieg, 1921; Billett et al., 1985).
This taxon is characterised by a flattened, elongated
body with a well developed lateral fringe (figure 76). It
measures a maximum of 20 cm long and 7 cm wide.
The lateral fringe has a dark tinge contrasting with the
A similar form, Benthodytes sanguinolenta (Théel, 1882),
from in the north-east Pacific Ocean, has also been found
in the Peruvian Basin, where it is relatively abundant. It
is an active swimmer and moves by rhythmic flexing.
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Fig. 75. Photograph taken by ‘Deep Tow’ at the Echo 1 site: nodule-facies C, holothurian Pannychia moseleyi (HOL 26) © Ifremer
Fig. 76. Photograph taken by the ‘Epaulard’ at the NIXO 45 site: nodule-facies B, holothurian Benthodytes typica
(HOL 27) © Ifremer
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HOL 28
This holothurian measures 17 to 25 cm long by 2 to 4 cm
wide (figures 77, 78). It appears flattened with the extremities rounded. In photographs taken by the “Nautile”,
it is pale violet with clearly darker shades along the podial fringe and at the anterior extremity, which is slightly
more developed. On the dorsal surface, two rows of long
dispersed papillae can be distinguished, turned down
towards the substratum. Just as in the preceding taxa,
no trail could be detected after its passage over ocean
floor covered in nodules. This taxon appears to prefer a
habitat with a relatively dense covering of nodules and
does not appear to be mesopelagic, never having been
seen in the water column.
Pawson & Foell (1983) identified Benthodytes incerta at
the DOMES C site. Kaufman & Smith (1997) and Lauerman et al. (1996) did not find it at their station further
to the north-east, and neither did Bluhm (1994; Bluhm
& Gebruk, 1999) in inventories of the Peruvian Basin. It
also appears to be absent from the Bay of Biscay in the
northern Atlantic Ocean (Sibuet et al., 1980).
The proposed identification, based on the literature (Ludwig, 1894; Oshima, 1915, 1915-1919; Clark, 1920; Madsen,
1953; Hansen, 1975) and on the advice of specialists, is
Benthodytes incerta (Ludwig, 1894), family Psychropotidae, Order Elasipodida. Benthodytes incerta has been
collected in the tropical east Pacific Ocean and around
Japan at depths varying from 2 417 m to 4 087 m (appendix 3). It has as external characters a semi-cylindrical body, a firm tegument finely covered with swellings,
9 to 15 tentacles, a narrow podial fringe delimiting the
well defined ventral sole, six to nine filiform or conical
dorsal papillae in two rows, 9-15 tentacles, circum-oral
papillae, a contractile peribuccal membrane, often turned down over the tentacles, and medio-ventral podia
arranged irregularly in a double row.
HOL 29
This dark taxon measures between 15 and 20 cm long
and 2-3 cm wide (figure 79). It has some long scattered
dorsal papillae extending over the dorsal radii. The peribuccal region is clearly distinct from the rest of the body.
It is never found in the water column but is present on
all types of facies with nodules and it leaves no visible
trail.
Pawson & Foell (1983) have identified three sorts of
Benthodytes from the DOMES C site, Benthodytes sp.,
B. incerta and B. typica, one of which is similar to HOL
29. Kaufman & Smith (1997) and Lauerman et al. (1996)
did not record Benthodytes sp. at their station. Bluhm
(1994) and Bluhm & Gebruk (1999) reported two types
of Benthodytes in their inventories from the Peruvian
Basin, as well as Benthodytes sanguinolenta, which is
very characteristic of this zone. This taxon was not recognised in photographs from the Bay of Biscay (Sibuet et
al., 1980).
We propose as a hypothesis of identification, based
on the literature (von Marenzeller, 1893; Ludwig, 1894;
Perrier, 1896; Clark, 1920; Deichmann, 1930, 1940, 1954;
Heding, 1940, 1942; Hansen, 1956, 1975) and the advice of specialists, a holothurian that is probably new
to science and related to the genus Benthodytes, B. sp.
type 2 (Théel, 1882), family Psychropotidae and Order
Elasipodida. One possibility would be Benthodytes lingua (Perrier, 1896), which is characterised by a podial
sole scarcely differentiated from the rest of the body,
filiform and conical dorsal papillae in up to 12 pairs,
and rounded ends to the body. Benthodytes lingua has
been collected in the Atlantic Ocean (04°05’N, 02°13’O)
at 2 100 m depth.
HOL 30
This holothurian measures about 30 cm long by 5 cm
wide (figures 77, 78). The tegument is off-white seems
randomly speckled by numerous small papillae. The
body appears cylindrical and rounded at the extremities. This taxon only appears on nodular ocean floor and
does not seem to be mesopelagic.
In referring to the literatures (Ludwig, 1893, 1894; Clark,
1920; Hansen, 1956) and on the advice of specialists,
we propose as a hypothesis of identification a species
of the genus Mesothuria (Théel, 1886), family Synallactidae, Order Aspidochirotida. Three members of this
genus have been collected not far from the study zone:
Mesothuria megapoda (Clark, 1920) at 4 245 m in the
central east Pacific Ocean, Mesothuria murrayi (Théel,
1886) which corresponds to HOL 9 described previously,
and Mesothuria multipes (Ludwig, 1893) from the Indian
Ocean and the central east Pacific Ocean between 725 m
and 4 064 m depth (appendix 3).
Pawson & Foell (1983) did not identify any Mesothuria
species from the DOMES C area except for M. murrayi,
and the same is true for Bluhm & Gebruk (1999) in the
Peruvian Basin.
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Fig.77. Photograph taken by the ‘Epaulard’ at the NIXO 45 site: nodule-facies C, holothurian Benthodytes incerta
(HOL 28) © Ifremer
Fig.78. Photograph taken by the ‘Nautile’
during the NIXONAUT cruise: facies of
plates, holothurian Benthodytes sp. type 1
(HOL 6) © Ifremer
Fig. 79. Photograph taken by ‘Deep
Tow’ at the Echo I site: mixed nodulefacies with plates (B and C), holothurian
Benthodytes sp. type 2 (HOL 29)
© Ifremer
Fig.80. Photograph taken by the ‘Epaulard’ at
the NIXO 45 site: nodule-facies C, holothurian
Mesothuria sp. (HOL 30)
© Ifremer
ENGLISH ATLAS PHOTO pages.indd 45
Fig.81. Photograph taken by the ‘Epaulard’ at the NIXO
45 site: nodule-facies C, holothurian Mesothuria sp. (HOL
30) © Ifremer
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HOL 31
These small holothurians 3-5 cm long and 1-2 cm wide
are difficult to see and can be found on all types of ocean
floor. They can sometimes be confused with other organisms such as sponges, molluscs or crustaceans. The
body is almost globular and bears some long dorsal
papillae and long, widely spaced, latero-ventral podia.
This taxon is only seen on sedimentary surfaces. The
small size of these holothurians is perhaps the reason
why they were not distinguished in photographs from
the Peruvian Basin.
Based on the literature (Clark, 1920; Hansen, 1956, 1967,
1975; Belyaev 1971; Lemche et al., 1976) and specialist
advice, identification proposed is a member of the family Elpidiidae, Order Elasipodida. Scotoplanes globosa (Théel, 1879), a cosmopolitan holothurian which
has been recorded in the study zone, is a possibility, or
Scotoplanes clarki (Hansen, 1975), which has only been
collected in the central eastern Pacific Ocean (10°N,
10°S) at depths between 3 570 m and 5 107 m (appendix 3). These holothurians both have two or three pairs
of long dorsal papillae, five to seven pairs of ventral
podia and 10 peribuccal tentacles. A smooth tegument
and rigid dorsal papillae would correspond to the external characters of Scotoplanes globosa, while a tegument covered with swellings and long, thin dorsal
papillae are characteristic of Scotoplanes clarki. These
holothurians can measure up to 9 cm. They have a very
special mode of locomotion, as mentioned previously
for HOL 10, using their long latero-ventral podia activated independently by the constriction of cavities
of the water-vascular system contained in the dermis
(Hansen, 1972). Some Scotoplanes species are known
to be occasional scavengers, feeding in fish carcasses
(Pawson, 1976).
The collected holothurians, which on the advice of specialists appear similar to the taxa identified in the Clarion-Clipperton fracture zone, are presented in appendix
3 with their geographic and bathymetric distribution.
46
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IOC TECHNICAL SERIES NO. 69, VOL.2
Appendix 1
List of codes
based on the classification of Parker (1982) (1982)
XEN
Xenophyophora, Cl. Xenophyophorea, Supercl. Rhizopoda, Phyl. Protozoa
SPO
Sponges, Phyl. Porifera
HYD
Hydroids, O. Hydroidea, Cl. Hydrozoa, Phyl. Cnidaria
MED
Jellyfish, O. Coronatae and O. Semaeostoniae, Cl. Scyphozoa, Phyl. Cnidaria
SIF
Siphonophores, O. Siphonophora, Cl. Hydrozoa, Phyl. Cnidaria
OCT
Octocoralliarians, O. Gorgonacea and O. Pennatulacea, Cl. Anthozoa, Phyl. Cnidaria
CER
Ceriantharians, O. Ceriantharia, Cl. Anthozoa, Phyl. Cnidaria
CTEN
Ctenophorans, Phyl. Ctenophora
ANT
Antipatharians, O. Antipatharia, Cl. Anthozoa, Phyl. Cnidaria
ACT
Sea-anemones, O. Actinaria, Cl. Anthozoa, Phyl. Cnidaria
CEP
Cephalopods, Cl. Cephalopoda, Phyl. Mollusca
NUD
Nudibranchs, O. Nudibranchia, Cl. Gastropoda, Phyl. Mollusca
MOL
Molluscs, Phyl. Mollusca
POL
Polychaete worms, Cl. Polychaeta, Phyl. Annelida
SIP
Sipunculians, Phyl. Sipuncula
ECH
Echiurians, Phyl. Echiuria
PYC
Pycnogonids, Cl. Pycnogonida, Phyl. Arthropoda
PER
Peracarids, Super-O. Peracaridea, Cl. Crustacea, Phyl. Arthropoda
ISO
Isopods, O. Isopoda, Super-O. Peracaridea, Cl. Crustacea, Phyl. Arthropoda
DEC
Decapods, O. Decapoda, Super-O. Peracaridea, Cl. Crustacea, Phyl. Arthropoda
GAL
Galatheas, Fam. Galatheidea, Infra-O. Anomourea, O. Decapoda, Super-O.
Peracaridea, Cl. Crustacea, Phyl. Arthropoda
CRI
Crinoids, Cl. Crinoïdea, Phyl. Echinodermata
AST
Seastars, Subcl. Somasteroïdea, Cl. Asteridae, Phyl. Echinodermata
OPH
Brittle stars, Subcl. Ophiuroidea, Cl. Asteridae, Phyl. Echinodermata
OUR
Sea urchins, Cl. Echinoidea, Phyl. Echinodermata
HOL
Sea cucumbers, Cl. Holothuridea, Phyl. Echinodermata
ENT
Epibenthic hemichordates, Cl. Enteropneusta, Phyl. Hemichordata
ASC
Ascidians, Cl. Ascidiacea, Subphyl. Tunicata, Phyl. Chordata
SAL
Salps, O. Salpidae, Cl. Thaliacea, Subphyl. Tuniciata, Phyl. Chordata
POI
Fish, Cl. Osteichthya, Subphyl. Vertebrata, Phyl. Chordata
47
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IOC TECHNICAL SERIES NO. 69, VOL.2
Appendix 2
List of international specialists consulted
Phylum PROTOZOA
Phylum PORIFERA
Phylum CNIDARIA
Cl. Xenophyophorea
U.K.
O. Kamenskaya
Federation of Russia
L. Levin
U.K.
C. Maybury
U.K.
O. Tendal
Denmark
Cl. Demospongiae
N. Boury-Esnault
Fr., Endume
Cl. Hexactinellida
W. Hartman
E.U.
V. Koltun
Federation of Russia
C. Levi
Fr., Muséum
D. Pawson
E.U.
O. Tendal
Denmark
J. Vacelet
Fr., Endume
Cl. Hydrozoa
O. Siphonophora
J. Bouillon
Belgium
J. Goy
Fr., Muséum
M. Van Praët
Fr., Muséum
W. Vervoort
Netherlands
C. Carré
Fr., Villefranche s/mer
F. Pagès
Spain
P. Pugh
U.K.
Cl. Scyphozoa
J. Bouillon
Belgium
Cl. Anthozoa
F.M. Bayer
E.U.
subcl. Alcyonaria
M. Grasshoff
Fed. Rep. of Germany
O. Gorgonacea
M.J. d’Hondt
Fr., Muséum
O. Pennatulacea
M. Opresko
E.U.
H. Zibrowius
(Fr., Endume)
subcl. Zoantharia
D. Doumenc
(Fr., Muséum)
O. Actinaria
D. Fautin
E.U.
K. Riemann-Zürneck
Fed. Rep. of Germany
M. Sokolova
Federation of Russia
O. Zezina
Federation of Russia
H. Zibrowius
Fr., Endume
O. Scleractinia
H. Zibrowius
Fr., Endume
O. Antipatharia
M. Grasshoff
Fed. Rep. of Germany
O. Ceriantharia
Phylum CTENOPHORA
A. Gooday
M.J. d’Hondt
Fr., Muséum
M. Opresko
E.U.
H. Zibrowius
Fr., Endume
D. Doumenc
Fr., Muséum
D. Fautin
E.U.
K. Riemann-Zürneck
Fed. Rep. of Germany
H. Zibrowius
Fr., Endume
C. Carré
Fr., Villefranche s/mer
48
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Phylum ANNELIDA
Cl. Polychaeta
Phylum ECHIURA
Phylum SIPUNCULA
Phylum MOLLUSCA
Cl. Bivalvia
Cl. Gastropoda
O. Nudibranchia
Cl. Cephalopoda
C. Bussau
Fed. Rep. of Germany
D. Desbruyères
Fr., Ifremer
L. Laubier
Fr., Ifremer
T. Miura
Japan
F. Pleijel
Sweden
E. Southward
U.K.
S. Edmonds
Australia
S. Ohta
Japan
C. Smith
E.U.
C. Bussau
Fed. Rep. of Germany
E. Cutler and N. Cutler
E.U.
P. Gibbs
U.K.
M. Rice
E.U.
P. Bouchet
Fr., Muséum
M. Rex
E.U.
A. Waren
Sweden
P. Bouchet
Fr., Muséum
M. Rex
E.U.
A. Waren
Sweden
C. Poizat
Fr., Marseille
R. Turner
E.U.
G. Aldred
U.K.
S. von Boletzky
Fr., Banyuls
M. Clarke
U.K.
T. Kubodera
Japan
M. NIXOn
U.K.
T. Okutani
Japan
C. Roper
E.U.
N. Voss
E.U.
R. Young
E.U.
C. Allan Child
E.U.
Phylum ARTHROPODA
Subph. CHELICERATA
Subph. CRUSTACEA
Cl. Pycnogonida
Super O. Peracarida
M. de Saint Laurent
Fr., Muséum
J. Stock
Netherlands
M. Türkay
Fed. Rep. of Germany
J. Svavarsson
Iceland
M. Türkay
Fed. Rep. of Germany
O. Mysidacea
J.P. Lagardère
Fr., Endume
O. Cumacea
M. Ledoyer
Fr., Endume
O. Tanaidacea
C. Allan Child
E.U.
A.B. Williams
E.U.
C. Allan Child
E.U.
R. Hessler
E.U.
J. Stromberg
Sweden
J. Svavarsson
Iceland
D. Thistle
E.U.
O. Isopoda
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IOC TECHNICAL SERIES NO. 69, VOL.2
O. Amphipoda
A.B. Williams
E.U.
G. Wilson
Australia
Ph. Laval
Fr., Villefranche s/mer
subo. Hyperiidea
O. Decapoda
C. Allan Child
E.U.
Natantia
A. Crosnier
Fr., Museum
J.P. Lagardère
Fr., Endume
M. Türkay
Fed. Rep. of Germany
A.B. Williams
E.U.
C. Allan Child
E.U.
A. Crosnier
Fr., Museum
I. Gordon
U.K.
J.P. Lagardère
Fr., Endume
M. de Saint Laurent
Fr., Museum
J. Stock
Netherlands
M. Türkay
Fed. Rep. of Germany
A.B. Williams
E.U.
A. Clark
U.K.
J. Durham
E.U.
D. Pawson
E.U.
Reptantia
Reptantia (suite)
Phylum ECHINODERMATA
Subph. CRINOZOA
Subph. ASTEROZOA
Subph. ECHINOZOA
Cl. Crinoidea
M. Roux
Fr., Reims
Cl. Stelleroidea
D. Blake
E.U.
subcl. Asteroidea
A. Clark
U.K.
L. Maluf
E.U.
D. Pawson
E.U.
M. Sibuet
Fr., Ifremer
subcl. Ophiuroidea
A. Clark
U.K.
O. Ophiurida
G. Hendler
E.U.
L. Maluf
E.U.
D. Pawson
E.U.
M. Sibuet
Fr., Ifremer
A. Gebruk
Federation of Russia
G. Hendler
E.U.
D. Pawson
E.U.
M. Roux
Fr., Reims
M. Sibuet
Fr., Ifremer
Cl. Echinoidea
Cl. Holothuroidea
Phylum HEMICHORDATA
Cl. Enteropneusta
D. Billett
U.K.
R.S. Carney
E.U.
A. Gebruk
Federation of Russia
M. Jangoux
Belgium
L. Maluf
E.U.
C. Massin
Belgium
D. Pawson
E.U.
M. Sibuet
Fr., Ifremer
M. Romero-Wetzel
Federation of Russia
E. Southward
U.K.
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Phylum CHORDATA
Subph. TUNICATA
Subph. VERTEBRATA
Cl. Ascidiacea
L. Cole
E.U.
C. Monniot
Fr., Museum
F. Monniot
Fr., Museum
Cl. Thaliacea
J.C. Braconnot
Fr., Villefranche s/mer
Cl. Osteichthyes
E. Anderson
South Africa
Cl. Mammalia
P. Castle
New Zealand
D. Cohen
E.U.
P. Geistdoerfer
Fr., Museum
T. Iwamoto
E.U.
N. Merrett
U.K.
J. Nielsen
Denmark
B. Seret
Fr., Museum
G. Demuizan
Peru
E. Hussenot
Fr., Brest
D. Robineau
Fr., Museum
51
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IOC TECHNICAL SERIES NO. 69, VOL.2
Appendix 3
Identification, geographic and bathymetric
distribution of echinoderms collected in the Pacific
Ocean
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF CRINOIDS COLLECTED
IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
52
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF CRINOIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
53
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION
AND GEOGRAPHIC AND BATHYMETRIC DISTRIBUTIONS OF ECHINOIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
54
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF OPHIUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
55
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IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF OPHIUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
56
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF ASTEROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
57
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF ASTEROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
58
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF HOLOTHUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
59
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF HOLOTHUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
60
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF HOLOTHUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
61
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IOC TECHNICAL SERIES NO. 69, VOL.2
IDENTIFICATION,
GEOGRAPHIC AND BATHYMETRIC DISTRIBUTION OF HOLOTHUROIDS
COLLECTED IN THE
Taxinomic
identification
Taxon
Authors + date
of identification
PACIFIC OCEAN
Gegraphic
distribution
Bathymetric
distribution
Substratum
Codes of taxons
on photographs and
specialists consulted
62
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Intergovernmental Oceanographic Commission
Technical Series 69
Biodiversity
and
distribution
of the
megafauna
Vol.2 Annotated photographic Atlas of the echinoderms
of the Clarion-Clipperton fracture zone
.......... . . . . . . . . .
Intergovernmental Oceanographic Commission (IOC)
United Nations Educational, Scientific and Cultural Organization (UNESCO)
1, rue Miollis,
75732 Paris Cedex 15,
France
Tel: +33 1 45 68 39 83
Fax: +33 1 45 68 58 12
Website: http://ioc.unesco.org
UNESCO
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